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Farkaë J. & Wrase D.W. - A new species of genus Leistus Frölich, 1799 from the Chinese Province 
of Gansu and new data on species previously described from Qinghai and Gansu (Coleoptera: 
Carabidae BNE brill) ocean ee une OS ON POP D EE D RO EE NT NE 


Serbina L., Burckhardt D. & Borodin O. - The jumping plant-lice (Hemiptera: Psylloidea) of 
BE] an SER nn ne cnrs rien baie Sate cossscen vats cecedunseuceesienadecuansaciedvevnesdandeancess 


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Kaydan M.B., Konczné Benedicty Z. & Szita E. - A new species of the genus Matileortheziola 
Kozancanoldil(Hemipteras Coccoideay Ontheziid ac) re 


Löbl I. - On the Scaphidiinae (Coleoptera: Staphylinidae) of the Lesser Sunda Islands.................. 


Lienhard C. & Ferreira R.L. - Review of Brazilian cave psocids of the families Psyllipsocidae 
and Prionoglarididae (Psocodea: ‘Psocoptera’: Trogiomorpha) with a key to the South American 
SPECIESIOMMESEIAMI LIES ere ottenere tds const 


Vogel P. & Gander A. - Live trapping design for the harvest mouse (Micromys minutus) in its 
SUIMMEMGNADILA E AI NOTTI 


de Chambrier A., Kuchta R. & Scholz T. - Tapeworms (Cestoda: Proteocephalidea) of teleost 
fishes from the Amazon River in Peru: additional records as an evidence of unexplored species 
CIVEFSINA RA RI NI IL I Lana 


Yin Z.-W. & Li L.-Z. - Review of the Himalayan genus Hingstoniella Jeannel, and description of 
Besuchetaceus gen. n. from Nepal (Coleoptera: Staphylinidae: Pselaphinae) ................................. 


7-44 
45-54 
55-70 


71-74 


75-120 


121-142 


143-148 


149-163 


165-180 


Revue suisse de Zoologie (March 2015) 122 (1): 1-5 


ISSN 0035-418 


A new species of genus Leistus Frölich, 1799 from the Chinese Province of Gansu 
and new data on species previously described from Qinghai and Gansu 
(Coleoptera: Carabidae: Nebriini) 


Jan Farkaë! & David W. Wrase? 


! Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague, Kamycka 1176, CZ-165 21 Prague 


6, Czech Republic. E-mail: farkac@fld.czu.cz 


> 


Dunckerstrasse 78, D-10437 Berlin, Germany. E-mail: carterus@gmx.de 


Abstract: A new species of genus Leistus Frölich, 1799, belonging to subgenus Evanoleistus Jedlicka, 1965 is described 
and illustrated: L. rezabkovae sp. n. from Gansu (type locality: Lenglong Ling Mts., Wutai Ridge [pass], 70 km N Honggu, 
3530 m, 36°58°16.6”N/102°48’03.6”E). It is compared with a similar species of the subgenus Evanoleistus, known from 
a mountain massif between the provinces Qinghai and Gansu. A check-list of all members of genus Leistus from the 
provinces Qinghai and Gansu is provided, comprising data on type locality and deposition of holotype. 


Keywords: Taxonomy - new distribution data - Palaearctic Region - China. 


INTRODUCTION 


Seven species of the genus Leistus Frölich, 1799, subgenus 
Evanoleistus Jedlitka, 1965, were recorded by Farkaë 
(2003, 2005) from Qinghai and Gansu Provinces (China). 
However, the two works do not include information about 
the occurrence of the species L. Jangmusianus in Gansu 
(Farkaë, 1999). Three species were described by Allegro 
(2007) from the southern part of the Gansu Province and 
finally two species were described by Deuve (2010) from 
Gansu. Based on new material collected in 2011, we are 
presenting new faunistic data and describing a species 
new to science. Thirteen species of the genus Leistus are 
thus currently known from the Gansu Province and only 
one species is known from the Qinghai Province (see 
below). 


MATERIAL AND METHODS 


Material examined is deposited in the collections of 

institutions and persons listed below: 

MNHUB Museum fiir Naturkunde der Humboldt- 
Universitat, Berlin, Germany 

MHNG Muséum d’Histoire Naturelle, Genève, 
Switzerland 

MNHN Muséum National Histoire Naturelle, Paris, 
France 

cFAR Coll. Jan Farkaë, Prague, Czech Republic 

cSCHM Coll. Joachim Schmidt, Admannshagen, 
Germany 

cWR Coll. David W. Wrase, Berlin, Germany 


Manuscript accepted 03.10.2014 
DOI: 10.5281/zenodo.14573 


Total body length (BL) is measured from the clypeus 
to the apex of the right elytron; the width of the head 
(HW) as the maximum linear distance across the head, 
including the compound eyes; the length of the pronotum 
(PL) between the anterior and the posterior margins 
along the midline; the length of the elytra (EL) from the 
tip of scutellum to the apex of the right elytron; the width 
of the pronotum (PW) and elytra (EW) at their broadest 
points; the width of the pronotal base (PBaW) between 
the tips of the hind angles. These measurements, made 
at a magnification of 16x and 32x by using an ocular 
micrometer in a stereobinocular microscope MBS10, 
were combined in ratios or added as follows: 

PW/PL: width/length of pronotum; PW/HW: width of 
pronotum/width of head; PW/PBaW: width of pronotum/ 
width of pronotal base; EL/EW: length/width of elytra; 
EW/PW: width of elytra/width of pronotum; EL/PL: 
length of elytra/length of pronotum. 

Line drawings were prepared using an ocular grid (15 x 
15 squares) attached to a Leica MZ 16 stereobinocular 
microscope. 

The photographs were taken with a Leica DFC450 digital 
camera (attached to Leica M205-C) using a motorised 
focussing drive, light base Leica TL5000 Ergo, diffused 
light with Leica hood LED5000 HDI, subsequently 
processed with Leica LAS application software, and 
enhanced with CorelDRAW Graphics Suite XS. 

Labels of type specimens are cited as in the original, the 
line break indicated by a forward slash (/). 


N 


J. Farkaë & D.W. Wrase 


TAXONOMY 


Leistus (Evanoleistus) rezabkovae sp. n. 
Figs 2-4 


Holotype: cFAR, without registration number; “China 
(Gansu Prov.) / Lenglong Ling Mts., Wutai Ridge / (pass), 
70 km N Honggu, 3530 m/36°58”16.6”N/102°48°03.6”E 
/ (stony alpine pasture with / Rhododendron and Azalea / 
under stones) / 2.VII.2011, D. W. Wrase [14]”, one male. 


Paratypes: cFAR; same locality and data as holotype, 
one male and one female. — cWR; same locality and 
data as holotype, two males. — cSCHM; same locality 
and data as holotype, but M. Schülke Igt., one male. — 
cFAR; “CHINA (Gansu Prov.) / Lenglong Ling Mts. / 
80 km NNW Honggu, 3392-3900 m / 37°03’50.3”N / 


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102°39°57.2”E / (alpine pasture with Rhododendron | 
under stones) / 30./31.VI.2011 D.W. Wrase [13]”, one 
male and one female. — cWR; same locality and data, one 
males and two females. - MHNG, without registration 
number; same locality and data, one female. - MNHUB; 
same locality and data, one female. — cWR; without 
registration number; the same data, but M. Schülke Igt., 


one male. — cWR; without registration number; same 
data but 3392 m, 1. VII.2011; one male. 


Diagnosis: Leistus rezabkovae sp. n. is similar in 
habitus to Leistus (Evanoleistus) farkaci Sciaky, 1994 
only, but the new species differs in the shape of elytra 
and the structure of the aedeagal median lobe. 

Morphometric data of male (holotype) (in mm): BL = 
7.90, HW = 1.61, PW = 1.93, PBaW = 1.03, PL = 1.50, 


(1) Leistus (E.) farkaci Sciaky, 1994. Habitus (Daban Shan, Pass 19 km WSW Men Yuan). 
(2) Leistus (E.) rezabkovae sp. n. Habitus (paratype, type locality). (Scale bar: 2 mm). 


New species of genus Leistus 3 


EW =2.90, EL = 4.62. Indices: PW/PL = 1.29, PW/HW = 
1.20, PW/PBaW = 1,87, EL/EW = 1,60, EW/PW = 1.50, 
EL/PL = 3.09. 

Morphometric data of 13 paratypes (in mm): 7 males: BL 
= 7.40-8.20, HW = 1.56-1.73, PW = 1.83-2.00, PBaW = 
0.90-1.03, PL = 1.40-1.56, EW = 2.75-2.98, EL = 4.50- 
4.75 and 6 females: BL = 8.00-8.50, HW = 1.63-1.70, 
PW = 1.90-2.00, PBaW = 0.98-1.05, PL = 1.50-1.63, EW 
= 2.88-3.05, EL = 4.63-5.00. Indexes. Males: PW/PL = 
1.23-1.33, PW/HW = 1.15-1.18, PW/PBaW = 1.83-2.03, 
EL/EW = 1.59-1.69, EW/PW = 1.45-1.53, EL/PL = 3.02- 
3.25. Females: PW/PL= 1.19-1.27, PW/HW = 1.13-1.20, 
PW/PBaW = 1.88-1.99, EL/EW = 1.58-1.66, EW/PW = 
1.48-1.55, EL/PL = 2.91-3.09. 


Description: Colour piceous-black, tarsi, tibiae, 
antennae and maxillary appendages paler, brownish. 
Mandibles brownish. Eyes prominent, convex. Gular 
setae not inserted on transverse carina (habitus of 
holotype Fig. 2). 

Pronotum: Cordiform, maximum width at middle, 
with distinct midline. Lateral furrow narrow, with 
one setiferous pore behind middle. Basal depression 
and depression at anterior margin strongly punctured. 
Posterior angles rectangular. 

Elytra: Long-oval, distinctly drop-shaped, maximum 
width somewhat behind middle, humeral angle 
completely reduced. Striae and lateral groove of elytra 
strongly punctured. Lateral groove in direction of elytral 
apex rather indistinct. 


Median lobe of aedeagus (Fig. 3, 4) relatively wide, 
about triangular apically, rounded at tip, apically with 
two chitinized ligaments laterally, consisting of granule- 
like structures aranged in a row (ventral view). In lateral 
view, the median lobe only gently curved toward apex. 


Comparisons: L. rezabkovae sp. n. differs from 
L. farkaci Sciaky by a different shape of elytra — 
L. rezabkovae sp. n. has elytra more distinctly drop- 
shaped , widest somewhat behind middle thirds (more 
or less oval elytra, compare Figs 1, 2), by pronotum 
shape, which is cordiform with rectangular posterior 
angles (in L. farkaci posterior angles obtuse-angled), by 
stronger punctuation of elytral striae (see Figs | and 2), 
and by a different shape of median lobe of aedeagus — 
in lateral view, L. farkaci has a median lobe narrower 
and apically somewhat more acuminate, distinctly bent 
toward apex (compare Figs 3-4 with 5, 6). 


Etymology: The new species is cordially dedicated to 
Mrs. Klara Rezäbkovä (Prague), a very good friend of 
the first author. 


Distribution: Currently known from two localities 
in the Lenglong Ling Mts. in the Chinese province of 
Gansu only. 


Habitat: Collected on a stony alpine pasture with 
Rhododendron sp. and Azalea sp. (Fig. 7) from under 
stones or by sifting litter and moss at an altitude of 
about 3392-3900 m. 


Figs 3-6. Leistus. Median lobe of aedeagus. (3) Leistus (E.) rezabkovae sp. n. (paratype, type locality), lateral view. (4) Leistus (E.) 
rezabkovae sp. n. (paratype, type locality), ventral view. (5) Leistus (E.) farkaci Sciaky (Daban Shan, Pass 19 km WSW Men 
Yuan), lateral view. (6) Leistus (E.) farkaci Sciaky (Daban Shan, Pass 19 km WSW Men Yuan), ventral view. (Scale bar: 


1 mm). 


Fig. 7. Photograph of type locality of Leistus (E.) rezabkovae sp. n. 


Leistus (Evanoleistus) farkaci Sciaky, 1994 


Leistus (Evanoleistus) farkaci Sciaky, 1994: 206-207. — Sciaky, 
1995: 300. — Farkaè, 1995: 159-160. — Farkaë, 1999: 41. — 
Farkaë, 2005: 49. 


New record: We examined 18 males and 6 females, 
labelled: China (Qinghai Prov.), Daban Shan, Pass 
19 km WSW Men Yuan, 3750-3900 m, 37°21’11.8”N 
/101°24°24.7"E, 4.VII.2011, D.W.Wrase Igt. [17] 
(cFAR, cSCHM, cWR, MNHUB). Specimens compared 
with paratypes in cFAR. Habitus Fig. 1. 


Habitat: Collected in an alpine cirque, on a stony 
pasture with slope springs in an altitude of 3750- 
3900 m. 


List of Leistus species (subgenus Evanoleistus Jedlitka, 
1967) from Gansu and Qinghai with data on the type 
locality (TL) and deposition of holotype (HT) 


Gansu 
becheti Allegro, 2007: 70 TL: “Lintan Xian, Yeliguan, 


J. Farkaë & D.W. Wrase 


Huan Hienzhi Natural Reserve” [HT in collection of 
Gianni Allegro] 

bohemorum Sciaky, 1994: 207 TL: “Golo Shan, Wen 
Xian” [HT in collection of Riccardo Sciaky, Milano] 

coltranei Allegro, 2007: 72 TL: “Weihuan Xian, Shimen 
Shan” [HT in collection of Gianni Allegro] 

gansuensis Sciaky, 1995: 296 TL: “between Xiahe and 
Heznojhen” [HT in collection of Riccardo Sciaky, 
Milano] 

heinzi Farkaë, 1995: 146 TL: “Da-li-Jia-Shan, 62 km W 
of Linxia” [HT in cFAR] 

huichuanensis Deuve, 2010: 3 TL: “montagnes a 24 km 
au sud-sud-est de Huichuan, 3700 m, 34°54’N/ 
104°04°E” [HT in MNHN] 

labrang Farkaë 1999: 24 TL: “Labrang, valley E of 
Ponggartang” [HT in collection of Miroslav Janata, 
Praha] 

langmusianus  Farkaë, 1995: 154 TL: Sichuan: 
“Langmusi” [HT in cFAR], data from Gansu (“Lugu, 
Dogcanglhamo”) in Farkaë, 1999: 39. 

lesteri Allegro, 2007: 69 TL: “Guan Shan, Caoyuan” [HT 
in collection of Gianni Allegro] 


New species of genus Leistus 5 


nanshanicus Belousov & Kabak, 2000: 105 TL: “Qilian 
Shan Mt. Range” [HT in collection of Zoological 
Institute of Russian Academy of Science, Sankt 
Petersburg] 

reflexus Semenov, 1889: 351 TL: “Amdo mts., Dzhoni” 
[HT in collection of Andreas Semenov Tian-Shanskij 
in Zoological Institute of Russian Academy of 
Science, Saint Petersburg] 

rezabkovae sp. n. TL: “Lenglong Ling Mts., Wutai 
Ridge” [HT in cFAR] 

xinglongensis Deuve, 2010: 4 TL: “monts Xinglong 
Shan, 8 km à l’ouest-sud-ouest de Mapo, 3500 m, 
35°46’N/103°55’E” [HT in MNHN] 

Qinghai 

farkaci Sciaky, 1994: 206 TL: “Daban Shan” [HT in 
collection of Museo Civico di Storia Naturale, 
Milano] 


ACKNOWLEDGEMENTS 


We thank our colleagues and friends who contributed 
to the realization of this paper, Miloslav Rakovic 
(Dobfichovice) for assisting in the first draft of this 
paper, Michael Schiilke for providing material this study 
deals with, Joachim Schmidt (Admanshagen) for making 
the habitus photographs, and Jon Cooter (Oxford) for 
reading a previous draft of the manuscript on which this 
paper is based. 


REFERENCES 


Allegro G. 2007. Three new Leistus species from Gansu (China) 
(Coleoptera Carabidae). Bollettino del Museo Civico di Sto- 
ria Naturale di Verona (Botanica Zoologia) 31: 69-73. 

Belousov I. A., Kabak I.I. 2000. A new species of the genus 
Leistus (Coleoptera: Carabidae) from China. Vestnik zoolo- 
gii 34(6): 105-108. 

Deuve T. 2010. Nouveaux Nebriidae, Broscidae et Trechidae de 
Chine et d’Iran (Coleoptera, Caraboidea). Revue Française 
d’Entomologie (N.S.) 32(1-2): 1-24. 

Farkaë J. 1995. Sixteen new species of Leistus from Asia 
(Coleoptera: Carabidae: Nebriini). Acta Societatis Zoologi- 
cae Bohemicae 59: 145-163 + 4 pp. 

Farkaë J. 1999. Check-list of the genus Zeistus (Coleoptera: 
Carabidae: Nebriini) from China with description of twenty- 
three new species. Folia Heyrovskyana, Supplementum 5: 
19-59. 

Farkaë J. 2005. Systematic Outline and Geographic Distribu- 
tion of Species of the Genus Leistus Frôlich, 1799 (Coleop- 
tera: Carabidae: Nebriini). Studies and Reports of District 
Museum Prague-East, Taxonomical Series 1(1-2): 43-67. 

Farkaë J., Janata M. 2003. Nebriini [pp. 18, 79-96]. /n: Löbl I. 
& Smetana A. (eds). The Catalogue of Palaearctic Coleop- 
tera. Vol. 1. Stenstrup: Apollo Books, 819 pp. 

Sciaky R. 1994. Seven new species of Leistus from China 
(Coleoptera: Carabidae: Nebriinae). Acta Societatis Zoolo- 
gicae Bohemicae 57 [1993]: 203-210. 

Sciaky R. 1995. Four new species of Leistus from China 
(Coleoptera, Carabidae, Nebriinae). Fragmenta Entomolo- 
gica 26(2): 293-303. 

Semenov A. P. 1889. Diagnoses coleopterorum novorum ex 
Asia centrali et orientali. Horae Societatis Entomologicae 
Rossicae 23: 348-403. 


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Revue suisse de Zoologie (mars 2015) 122 (1): 7-44 


ISSN 0035-418 


The jumping plant-lice (Hemiptera: Psylloidea) of Belarus 


Liliya Serbina':*, Daniel Burckhardt! & Oleg Borodin? 


!  Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Switzerland 
2? Institut für Natur-, Landschafts- und Umweltschutz an der Universität Basel, St. Johanns-Vorstadt 10, CH-4056 Basel, 


Switzerland 


3 National Academy of Sciences, Akademicheskaya street 27, 220072 Minsk, Belarus 
Corresponding author: Liliya Serbina. E-mail: liliia.serbina@unibas.ch, liliya_serbina@mail.ru 


Abstract: A checklist of the known psyllids of Belarus is given: 12 species (one of them doubtful) have been previously 
reported and 43 species are added here, bringing the number of confirmed species to 54. The psylloid fauna of the coun- 
try remains poorly known. Based on information from surrounding countries, another 73 species can be expected. An 
illustrated identification key is provided for the 127 species whose occurence in Belarus has been confirmed or is likely. 


Keywords: Psyllids - Sternorrhyncha - host-plants - faunistics - identification key - Europe - Palaearctic Region. 


INTRODUCTION 


Jumping plant-lice or Psylloidea, a superfamily of Ster- 
norrhyncha, are plant-sap sucking insects (Photo 1). 
Most psyllid species are associated with dicotyledonous 
plants, and are usually highly host-plant specific. At 
present there are almost 4000 described psyllid species 
world-wide (Li, 2011) representing probably less than 
half of the actually existing species number. Despite the 
fact that psyllids are most species-rich in the tropics and 
south temperate regions, the west Palaearctic fauna is 
the one which is best studied with around 400 species 
reported from Europe (Burckhardt, 2004). The study of 
Eastern European psyllids was initiated by Flor (1861) 
with a survey on the fauna of Livonia followed by a large 
number of publications dealing with the European part 
of the former Soviet Union (Gegechkori & Loginova, 
1990). 

The information on the psyllid fauna of Belarus is, how- 
ever, scarce with only 12 recorded species of which 
one is doubtful (Loginova, 1961, 1962b; Palyakova, 
1969; Byazdzenka et al., 1973; Gorlenko et al., 1988; 
Sidlyarevich & Bolotnikova, 1992; Petrov, 2004, 2011; 
Petrov et al., 2011; Petrov & Sautkin, 2013). 

In the last 15 years psyllids were collected in all admi- 
nistrative regions of Belarus, mostly as by-catch of an 
Auchenorrhyncha survey. The aim of the present paper is 
to provide an updated checklist of psyllids from Belarus 
based on literature data and recently collected material. 
We also provide an illustrated identification key for 
the species recorded and those potentially occurring in 
Belarus, as the existing keys are outdated or incomplete. 


Manuscript accepted 28.07.2014 
DOI: 10.5281/zenodo.14578 


MATERIAL AND METHODS 


Unless stated otherwise, the material was collected by 
O. Borodin, is dry mounted and deposited in the Labo- 
ratory of Terrestrial Invertebrates of the State Science 
and Production Association “Research and Practical 
Centre of the National Academy of Sciences of Belarus 
for Bioresources” (Gosudarstvennoye Nauchno-Proiz- 
vodstvennoye Ob‘yedineniye “Nauchno-Prakticheskiy 
Tsentr Natsional‘noy Akademii Nauk Belarusi po Biore- 
sursam”) in Minsk. Material from the Naturhistorisches 
Museum, Basel (Switzerland) was examined for compa- 
rison. 

For the transliteration of names and organisations from 
the Cyrillic to the Latin alphabet the BGN/PCGN system 
was used. 

Morphological terminology mainly follows Ossiannilsson 
(1992), Hollis (2004) (wing venation) and Burckhardt 
(2010). Botanical nomenclature and classification 
follows Sautkina e? al. (1999). Additional information on 
psyllid host-plants is available in Ossiannilsson (1992) 
and Ouvrard (2014). Generally useful identification 
keys are by Loginova (1964), Klimaszewski (1975) 
and Ossiannilsson (1992), though they are more or less 
outdated and should be used with caution. Photos and 
drawings were prepared from slide mounted specimens 
located in the Naturhistorisches Museum, Basel. 
Photos were made with KEYENCE VHX-2000 digital 
microscope. 


L. Serbina, D. Burckhardt & O. Borodin 


Photo 1. Living Livia junci, photo kindly provided by Gernot Kunz. 


Following abbreviations are used: 

Checklist: BSU — Belarusian State University, MKAD — 
Minsk automobile ring road (Minskaya kol‘tsevaya avto- 
mobil‘naya doroga). 

Identification key: AEL — length of distal portion of 
aedeagus, ALHW — antenna length : head width ratio, 
FPHW - female proctiger length : head width ratio, MP 
— male proctiger length. 


SYSTEMATIC ACCOUNT 


Checklist 
The checklist is alphabetical using the classification of 
Burckhardt & Ouvrard (2012). 


Aphalaridae 
Aphalarinae 
Aphalara affinis (Zetterstedt, 1828) 


Material examined: Minsk: | adult without abdomen, 
Molodechno district, nr Myasota, 6.1x.2002. 


Aphalara avicularis Ossiannilsson, 1981 


Material examined: Brest: 1 ©, Stolin district, Belousha, 
15.v111.2005. 


Comments: The record of A. polygoni by Loginova 
(1961) from the Vitebsk region may concern A. avicu- 
laris or A. freji. Without revising the original material it is 
not possible to decide which species is concerned. 


Aphalara freji Burckhardt & Lauterer, 1997 


Material examined: Gomel’: 14, Lel’chitsy district, 
0.5 km NE Krasnoberezh’ye, river Ubort’ floodplain, 
5.viii.2004. — Minsk: 19, Minsk district, Shchemyslitsy, 
Dubrava Natural Monument, 26.ix.2013, on conifers 
(L. Serbina). — 34, same but nr Aksakovshchina, Wild- 
life sanctuary Podsady, 28.ix.2013, mixed forest. — 54, 
19, same but nr Tarasovo, nr Gardeners’ partnership 
Zvezdnoye, 29.1x.2013. 


Comments: The record of 4. polygoni by Loginova 
(1961) from the Vitebsk region may concern A. avicula- 
ris or A. freji. Without revising the original material it is 
not possible to decide which species is concerned. 


Jumping plant-lice of Belarus 9 


Craspedolepta crispati Lauterer & Burckhardt, 2004 


Material examined: Minsk: 44, 27, Volozhin district, 
Rakov, 28.vii.1997, meadow. 


Craspedolepta flavipennis (Foerster, 1848) 


Material examined: Minsk: 19, Myadel’ district, 
Logoviny, train station, 5.vii.2009, meadow (L. 
Serbina). 


Craspedolepta latior Wagner, 1944 


Material examined: Minsk: 24, 19, Minsk district, 
river Ptich’ bank, nr train station Ptich’, 15.vii.2009 
(L. Serbina). — 19, same but Molodechno district, nr 
Vyazynka, canal bank, 9.vii.2002. — 114, 272, same but 
Myadel’ district, Logoviny, nr train station, 5.v11.2009, 
meadow (L. Serbina). — 14, 19, same but nr Naroch’, 
9.vii.2009. 


Craspedolepta malachitica (Dahlbom, 1851) 


Material examined: Gomel’: 19, Kalinkovichi dis- 
trict, 0.5 km E Turovichi, 23.vii.2005, dry meadow, 
on Artemisia sp. — Grodno: 1 adult without abdomen, 
Mosty district, 3.5 km NW Peski, river Zel’vyanka 
floodplain, 24.vi.2005 (Y. Gerashchenko). — Minsk: 
49, Minsk district, Shchemyslitsy, Dubrava Natural 
Monument, 15.v.2008 (L. Serbina). — 29, same but river 
Ptich’ bank, nr train station Ptich’, June 2008. — 19, 
same but Molodechno district, nr Povyazyn’, 1.v11.2001. 
— 25, same but nr canal bank, 9.vi.2002. — 14, same but 
nr Radoshkovichi, slope Minsk-Vileyka channel system, 
22.vi.2005. — 19, same but nr Sitsevichi, 12.v11.2005, 
pine forest (A. Egiyan). — 24, 69, same but river 
Udranka bank, nr bus stop, 14.vii.2009 (L. Serbina). 
— 14, 29, same but Myadel’ district, nr Naroch’, 
2.vii.2009. — 104, 249, same but Logoviny, nr train 
station, 5.vii.2009. — 844, 999, same but nr Naroch’, 
9.vii.2009. — 19, same but Volozhin district, nr Kaldyki, 
11.vii.2007, mixed forest. — Vitebsk: 29, Lepel’ dis- 
trict, nr Domzheritsy, nr boarding school and pond, 
31.vii.2001, on Artemisia dracunculus. — 19, same 
but river Buzyanka, 300 m upstream from boat station, 
1.viii.2001. 


Craspedolepta nervosa (Foerster, 1848) 
Material examined: Grodno: 19, Mosty district, 3.5 km 
NW Peski, 24.vi.2005, river Zel’vyanka floodplain. — 
Minsk: 1, Borisov district, 2 km NW Peresady, nr train 
station “Proletarskaya pobeda”, lowland, 6.vi11.2002, 
pine forest. — 19, same but Logoysk district, nr Besyady, 
river Udra floodplain, 26.vi.2002. — 19, same but Kupa- 


lovskiy National Park, nr Lekarovka, 7.vii.2004, dry 
meadow. — 4%, 39, same but 0.5 km SW Matski, hill, 
meadow. — 19, same but Minsk district, nr train station 
Kryzhovka, 3.vi.2001, meadow in pine forest. — 14, 19, 
same but nr Chirovichi, 9.vi.2002, dry meadow. — 14, 
same but Shchemyslitsy, nr Biological Faculty, BSU, 
18.vi.2002. — 19, same but Dubrava Natural Monu- 
ment, 19.vi.2002. — 19, same but nr Biological Faculty, 
BSU, 1.vii.2002. — 14, 32, same but 19.vi.2006 (L. Ser- 
bina). — 14, same but Molodechno district, nr Vyazynka, 
16.vi.2001, dry meadow. — 19, same but 2.vii.2001. 
— 14, same but 0.5 km E Vyazynka, hill, 3.vii.2001. — 
14, same but nr Vyazynka, canal bank, 12.vi.2002. — 
14, 39, nr Shipulichi, river Zapadnaya Berezina bank, 
27.vi.2002. — 14, same but river Udranka bank, nr bus 
stop, 14.vii.2009 (L. Serbina). — 14, 19, same but Mya- 
del’ district, nr Naroch’, 2.vii.2009. — 2°, 39, same but 
Logoviny, nr train station, 5.vii.2009, meadow. — 34: 
same but 0.5 km S Nikol’tsy, Gomza Dendrological Gar- 
den, 6.vii.2009. — 14, 19, same but Soligorsk district, 
7 km from Soligorsk, 24.vi.2004 (A. Egiyan). — 19, 
same but 25.vi.2004. — 19, same but Stolbtsy district, nr 
train station Kolosovo, 26.v.1993, field in front of pine 
forest (V. Karasev). — Vitebsk: 24, 29, Gorodok district, 
17.vi.2008, dry meadow, on Achillea millefolium. 


Craspedolepta omissa Wagner, 1944 


Material examined: Minsk: 19, Logoysk district, 
Kupalovskiy National Park, nr Lekarovka, 7.vii.2004, 
dry meadow. — 19, same but Minsk district, 2 km NE 
Shchemyslitsy, along train station, 12.vii.2005, forest 
plantation. — 24, 19, same but river Ptich’ bank, nr 
train station Ptich’, 15.vii.2009 (L. Serbina). — 2°, 
same but Botanical Garden (Minsk), 16.v11.2009. — 
14, same but Molodechno district, nr Vyazynka, sta- 
tion, hill, 3.vii.2001. — 24, 29, same but nr station 
Shipulichi, 27.vi.2002, river Zapadnaya Berezina bank, 
on Artemisia sp. — 16, same but Myadel’ district, nr 
Naroch’, 2.vii.2009 (L. Serbina). — 65, 109, same but 
Logoviny, nr train station, 5.vii.2009. — 1 adult without 
abdomen, same but nr Antonisberg, 300 m behind res- 
ting place, meadow. — 34, 59, same but 0.5 km S 
Nikol’tsy, Gomza Dendrological Garden, 6.vii.2009. 
— 19, same but nr Naroch’, 9.vii.2009. — 14, same 
but Vileyka district, nr hydraulic station, 0.5 km N 
Budishche, 9.vii.2005, dry meadow. 


Craspedolepta subpunctata (Foerster, 1848) 


Material examined: Minsk: 19, Molodechno district, 
nr Vyazynka, 16.vi.2001, willow thicket. — 19, same but 
nr Chirovichi, 9.v1.2002. 


10 L. Serbina, D. Burckhardt & O. Borodin 


Rhinocolinae 
Rhinocola aceris (Linnaeus, 1758) 


Material examined: No locality data, 14, 19 (S. 
Buga). 


Liviidae 
Euphyllurinae 
Psyllopsis discrepans (Flor, 1861) 


Material examined: Gomel’: 19, Khoyniki district, 
Orevichi, Polesskiy Natural Reserve, 21.viii.1991, 
Malaise trap. — Minsk: 154, 309, Soligorsk district, 
7 km from Soligorsk, 24.vi.2004 (A. Egiyan). 


Psyllopsis distinguenda Edwards, 1913 


Material examined: Vitebsk: 14, 19, Gorodok district, 
Carla-Marksa Street, 17.vi.2008, on Fraxinus sp. (S. 
Buga). 


Psyllopsis fraxini (Linnaeus, 1758) 


Records: Recorded by Petrov (2011) from the south 
central region of the Belarusian Ridge. 


Psyllopsis fraxinicola (Foerster, 1848) 


Material examined: Minsk: 14, Soligorsk district, 
7 km from Soligorsk, 24.v1.2004 (A. Egiyan). 


Strophingia ericae (Curtis, 1835) 


Material examined: Grodno: 2%, Iv’yev dis- 
trict, Naliboki pushcha, 4 km NE Malaya Chapun’, 
22.v1.2002, pine forest, on Calluna sp. 


Liviinae 
Camarotoscena speciosa (Flor, 1861) 
Material examined: Minsk: 14, Minsk, alley 
nr Lyubimov Ave., 23.iv.2013, on Tilia sp. — 39, 
same but Botanical Garden (Minsk), 26.iv.2013, on 
Rhododendron sp. (L. Serbina). 


Psyllidae 
Psyllinae 
Arytaina genistae (Latreille, 1804) 


Material examined: Vitebsk: 24, 19, Lepel’ district, 
nr Kraytsy, 1 km along road in direction Perechodtsy, 
2.viii.2001, pine forest. 


Baeopelma foersteri (Flor, 1861) 


Records: Recorded by Loginova (1962b) as Psylla 
foersteri from the Vitebsk region. 


Material examined: Gomel’: 24, 39, Zhitkovichi 
district, Khvoyensk, 11.vii.1999, pine forest, on A/nus 
glutinosa (S. Buga). — 34, same but 1.5 km SW Novyye 
Milevichi, river Sluch’ floodplain, 4.viii.2004. — 154, 
9°, same but 2 km SW Otskovannoye, 5.viii.2004. — 
14, same but nr Novyye Milevichi, nr river Sluch’, 
15.ix.2004. — Minsk: 14, 29, Logoysk district, 0.5 km 
SW Matski, hill, 7.vii.2004, meadow. — 19, same 
but Minsk district, Kupalovskiy National Park, nr 
Lekarovka, dry meadow, on Alnus incana. — 14, 1° 
same but Botanical Garden (Minsk), 16.vii.2009, on 
Alnus sp. (L. Serbina). — 24, 19, same but river Ptich’ 
bank, nr train station Ptich’, 22.vii.2009. — 34, 59, 
same but Myadel’ district, Naroch’, nr Antonisberg, 
30.vi.2005, (A. Egiyan). — 44, 39, same but 1.vii.2005, 
on Alnus incana. — 19, same but nr Naroch’, 3.vii.2009, 
on Alnus sp. (L. Serbina). — 14, same but 0.5 km S 
Nikol’tsy, Gomza Dendrological Garden, 6.vii.2009. — 
19, same but Soligorsk district, Soligorsk, 25.vi.2004 
(A. Egiyan). — 54, 49, same but Volozhin district, 
Rakov, river Isloch’ floodplain, 28.vii.2004. — 34, 69, 
same but on Alnus glutinosa. — 24, 29, same but nr 
Kaldyki, 19.vii.2007, mixed forest, on Alnus sp. (L. 
Serbina). — Mogilev: 19, Klichev district, Lozovitsa, 
20.vii.1997, pine forest. — Vitebsk: 19, Postavy district, 
lake Chetvert’ south bank, 10.vii.2005. 


Cacopsylla ambigua (Foerster, 1848) 


Material examined: Brest: 24, Drogichin dis- 
trict, Wildlife sanctuary “Zvonets”, 16.vi.1999. — 
Gomel’: 29, Zhitkovichi district, 1.5 km SW Novyye 
Milevichi, river Sluch’ floodplain, 4.viii.2004, on 
Salix sp. — Grodno: 1 adult without abdomen, Iv’yev 
district, Naliboki pushcha, 0.5 km S Potashnya, nr 
river, 22.v111.2002, meadow (E. Shestakov). — Minsk: 
44, 89, Logoysk district, nr Gayany, 28.vi.2004, on 
Salix sp. — 14, 29, same but Minsk district, train sta- 
tion Kryzhovka, river Poplav bank, 3.vi.2001, flood- 
plain meadow. — 29, same but nr Shchemyslitsy, NW 
Dubrava Natural Monument, 6.vi.2002. — 19, 1 adult 
without abdomen, same but 15.vi.2002. — 42, same but 
18.vi.2002. — 22, same but 4.vii.2002. — 14, 19, same 
but field between MKAD Kurasovshchina and South- 
West district, 24.v.2003. — 14, same but Shchemyslitsy, 
nr Biological Faculty, BSU, 19.vi.2006. — 14, 42, same 
but Molodechno district, nr Vyazynka, 16.vi.2001, 
bog. — 14, 39, same but nr station Shipulichi, river 
Zapadnaya Berezina floodplain, meadow edge, 
27.vi.2002. — 19, same but nr Vyazynka, 20.v.2003. — 
Vitebsk: 14, Gorodok district, 2 km NE Machalovo, 
river Lovat’ floodplain, 4.vi.2000, bog (S. Buga). 


Jumping plant-lice of Belarus Il 


Cacopsylla brunneipennis (Edwards, 1896) 


Material examined: Minsk: 24, 39, Logoysk district, 
Gayany, 28.vi.2004. — 24, 39, same but Minsk dis- 
trict, nr Aksakovshchina, Wildlife sanctuary Podsady, 
28.1x.2013, mixed forest, on conifers (L. Serbina). 


Cacopsylla crataegi (Schrank, 1801) 


Records: Recorded from Belarus by Gorlenko er al. 
(1988) as Psylla crataegi and by Petrov & Sautkin 
(2013) as Cacopsylla crataegi from the Minsk region. 


Cacopsylla hippophaes (Foerster, 1848) 


Material examined: Minsk: 14, 32, Minsk district, 
nr Tarasovo, nr Gardeners’ partnership Zvezdnoye, 
29.1x.2013, on Hippophae rhamnoides (L. Serbina). 


Cacopsylla ledi (Flor, 1861) 


Material examined: Gomel’: 19, Zhitkovichi dictrict, 
Pripyatskiy National Park, 11.vii.1999, pine forest, on 
Ledum sp. — Minsk: 19, Myadel’ district, 1 km SW 
Kochergi, 16.vii.2005, pine forest, on Ledum sp. — 19, 
same but Leytsy, 5.v.2008. — Mogilev: 19, Klichev dis- 
trict, Razvadovo, 20.vii.1997, pine forest, on Ledum 
sp. — 14, 19, same but Lozovitsa, 31.viii.1997. — 19, 
Razvadovo, 23.vii.1998. — 39, same but 17.viii.1998. 
— 14, same but, 17.ix.2000, pine forest. — Vitebsk: 
14, Dokshitsy district, Krulevshizna, 11.vii.2000, 
pine forest (L. Chumakov). — 19, same but 22.x.2000. 
— 19, same but Rossony district, 0.8 km N Osinniki, 
11.1x.1998, pine forest, on Ledum sp. 


Cacopsylla mali (Schmidberger, 1836) 


Records: Recorded by Byazdzenka (1973) as Psylla 
mali from the Minsk region. 


Material examined: Minsk: 14, Logoysk dis- 
trict, Kupalovskiy National Park, nr Lekarovka, 
7.vii.2004, dry meadow. — 14, 29, same but Minsk 
district, Shchemyslitsy, Dubrava Natural Monument, 
16.1x.1997, on Malus sylvestris (S. Buga). — 19, same 
but 15.vii.2009, on Malus sp. (L. Serbina). — 19, same 
but Molodechno district, nr Vyazynka, 16.vi.2001, bog. 
— 29, same but 29.vii.2001. — 19, same but nr sta- 
tion Shipulichi, river Zapadnaya Berezina floodplain, 
27.vi.2002. — 29, same but Myadel’ district, Naroch’ 
orchards, 2.vii.2009 (L. Serbina). — 19, same but 
Volozhin district, nr Kaldyki, 11.vii.2007, mixed forest. 


Cacopsylla mali (Schmidberger, 1836) group 


Material examined: Minsk: 19, Logoysk district, 
Kupalovskiy National Park, nr Lekarovka, 7.vii.2004, 
meadow. — 17, same but Minsk district, Shchemyslitsy, 
Dubrava Natural Monument, 15.vi.2002. — 19, same 
but nr Biological Faculty, BSU, 1.vii.2002. — 19, same 
but Molodechno district, nr Vyazynka, 1.vi.2001. — 1 
adult without abdomen, same but Myadel’ district, 
nr Naroch’, 4.vii.2009 (L. Serbina). — 14, same but 
Nesvizh district, nr Nesvizh, 29.vi.2004, on Prunus 
cerasifera. 


Comment: Due to the poor state of the material the 
specimens cannot be identified to species. They could be 
any member of the C. mali group: C. mali, C. peregrina, 
C. sorbi or C. ulmi. 


Cacopsylla moscovita (Andrianova, 1948) 


Material examined: Grodno: 14, 19, Iv’yev dis- 
trict, Naliboki pushcha, 4.5 km E Malaya Chapun’, 
22.v111.2002, on Salix sp. 


Cacopsylla parvipennis (Lòw, 1877) 


Material examined: Minsk: 19, Myadel’ district, 
2 km W Cheremshitsy, river Narochanka floodplain, 


9.v11.2005. 


Cacopsylla peregrina (Foerster, 1848) 


Material examined: Minsk: 34, 72, Logoysk dis- 
trict, Kupalovskiy National Park, nr Lekarovka, 
7.vii.2004, dry meadow. — 14, 19, same but Minsk dis- 
trict, stop “Kurasovshchina”, 9.ix.1997, on Crataegus 


sp. (S. Buga). — 49, same but stop “Bol’nitsa”, 
13.ix.1997. — 4%, 49, same but Shchemyslitsy, 
Dubrava Natural Monument, 21.v.1999. — 3%, 


same but station “Polzhelishche”, 18.ix.1999. — 29, 
same but Shchemyslitsy, track nr bus stop “Filial 
BGU”, 11.vii.2009 (L. Serbina). — 14, 19, same but 
22.vii.2009. — 159, same but 29.vii.2009. — 34, 49, 
same but Volozhin district, nr Kaldyki, 19.v11.2007, 
mixed forest. 


Cacopsylla pulchella (Löw, 1877) 


Material examined: Gomel’: 8%, 29, Khoyniki 
district, Orevichi, 21.v.-18.vi.1991, Malaise trap. — 24, 
same but Zhitkovichi district, 14 km from Chvoensk, 
Pripyatskiy National Park, 26.viii.1999, pine forest. 
— Minsk: 24, Molodechno district, nr Vyazynka, 
16.vi.2001, dry meadow. 


12 L. Serbina, D. Burckhardt & O. Borodin 


Cacopsylla pulchra (Zetterstedt, 1838) 
Material examined: Gomel’: 14, Zhitkovichi dis- 
trict, Pripyatskiy National Park, 20.x.1998, pine forest. 
— Minsk: 14, Logoysk district, Gayany, 28.vi.2005. — 
24, 12, same but Minsk district, nr Aksakovshchina, 
Wildlife sanctuary Podsady, 28.1x.2013, mixed forest, 
on conifers (L. Serbina). — 14, same but Molodechno 
district, nr Vyazynka, 16.v1.2001, bog, on Salix sp. 


Cacopsylla pyri (Linnaeus, 1758) 


Records: Recorded from Belarus by Palyakova (1969), 
Gorlenko er al. (1988) as Psylla pyri. 


Cacopsylla pyrisuga (Foerster, 1848) 


Records: Recorded from Belarus by Palyakova (1969) 
as Psylla pyrisuga. 


Cacopsylla saliceti (Foerster, 1848) group 


Material examined: Grodno: 19, Iv’yev district, 
Naliboki pushcha, 2 km S Potashnya, 23.vi.2002, on 
Salix sp. — 19, same but 4.5 km E Malaya Chapun’, 
28.ix.2002. — Minsk: 19, Minsk district, nr Kryzhovka 
station, 3.vi.2001, alder forest. — 19, same but Myadel’ 
district, Leytsy, 25.vi.2008. — 19, same but 0.5 km S 
Nikol’tsy, Gomza Dendrological Garden, 6.vii.2009, on 
Salix sp. (L. Serbina). 


Comment: Single females of this species group cannot 
be correctly identified to species. 


Cacopsylla sorbi (Linnaeus, 1767) 


Material examined: Minsk: 384, 329, Myadel’ dis- 
trict, Naroch’, nr Antonisberg, 28.vi.2005, on Sorbus sp. 
(A. Egiyan). 


Cacopsylla ulmi (Foerster, 1848) 


Records: Recorded by Petrov (2011) from the south 
central region of the Belarusian Ridge. 


Material examined: Gomel’: 34, 39, Khoyniki dis- 
trict, Orevichi, Pripyatskiy National Park, 23.vii.1991, 
Malaise trap. — 6¢, 329, same but Polesskiy Natural 
Reserve, 21.viii.1991. — 19, same but Zhitkovichi dis- 
trict, Khvoyensk, truck patch, 14.vii.1999 (S. Buga). 
— Minsk: 14, Minsk district, Shchemyslitsy, nr train sta- 
tion “Roshcha”, 15.vii.2009, on Ulmus sp. (L. Serbina). 


Chamaepsylla hartigii (Flor, 1861) 


Material examined: Minsk: 19, Logoysk district, 
Kupalovskiy National Park, nr Lekarovka, 7.vii.2004, 
dry meadow, on Betula pendula. 


Psylla alni (Linnaeus, 1758) 


Material examined: Brest: 45, 29, Gantsevichi dis- 
trict, 3 km SE Borki, 23.vii.2005, pine forest. - Gomel’: 
29, Zhitkovichi district, 1.5 km SW Novyye Milevichi, 
river Sluch’ floodplain, 4.viii.2004. — 34, 59, 2 km, 
same but SW Otskovannoye, 5.v111.2004. — Grodno: 
19, Mosty district, 3.5 km NW Peski, river Zel’vyanka 
floodplain, 23.vi.2005. — 19, same but 24.vi.2005. — 
Minsk: 624, 669, Logoysk district, nr Gayany, roadside 
lowland, 28.vi.2004, on Alnus incana. — 19, same but 
Kupalovskiy National Park, nr Lekarovka, 7.vi1.2004. 
— 34, 62, same but 0.5 km SW Matski, hill, meadow. 
— 34, 49, same but Molodechno district, nr Vyazynka, 
16.vi.2001, bog. — 64, 39, same but 1.vii.2001, on 
Alnus sp. — 24, 49, same but river Udranka bank, nr 
bus stop, 2.vii.2008, on Alnus incana (L. Serbina). — 
14, same but nr Naroch’, 3.vii.2009, on Alnus sp. — 29, 
same but Nesvizh district, park “Alba”, 29.vi.2004. — 
24, 52, same but Volozhin district, Rakov, river Isloch’ 
floodplain, 28.vii.2004, on Alnus glutinosa. — Vitebsk: 
14, Gorodok district, 0.3 km SW Zadrach’ye, river 
Zadrach bank, 7.vi.2000, on Alnus incana (S. Buga). 


Psylla betulae (Linnaeus, 1758) 


Material examined: Mogilev: 19, Klichev district, 
Razvadovo, 23.vii.1988, pine forest. 


Psylla buxi (Linnaeus, 1758) 


Records: Recorded by Petrov ef al. (2011), Petrov & 
Sautkin (2013) from the Minsk region. 


Psylla fusca (Zetterstedt, 1828) 


Material examined: Minsk: 19, Logoysk district, 
Kupalovskiy National Park, nr Lekarovka, 26.v1.2002, 
dry meadow. — 284, 239, same but nr Gayany, down- 
hill, 28.vi.2004, on Alnus incana. — 14, 19, same but 
Kupalovskiy National Park, nr Lekarovka, 7.vii.2004, 
dry meadow. — 54, 49, same but 0.5 km SW Matski, 
hill, meadow. — 1 adult without abdomen, same but 
Molodechno district, nr Vyazynka, station, 1.v11.2001. 
— 14, same but 2.vii.2001, on Alnus sp. — 1 adult with- 
out abdomen, same but 29.vii.2001, linden alley. — 
203, 172, same but river Udranka bank, nr bus stop, 
2.vii.2008, on Alnus incana (L. Serbina). — 14, same 


Jumping plant-lice of Belarus 13 


but Volozhin district, nr Kaldyki, 19.vii.2007, mixed 
forest, on Alnus sp. 


Triozidae 
Bactericera acutipennis (Zetterstedt, 1828) 


Material examined: Vitebsk: 34, 19, Dokshitsy dis- 
trict, nr station Krulevshizna, 20.x.2000, pine forest (L. 
Chumakov). 


Bactericera curvatinervis (Foerster, 1848) 


Material examined: Minsk: 14, Minsk district, 
nr Aksakovshchina, Wildlife sanctuary Podsady, 
28.ix.2013, mixed forest, on conifers (L. Serbina). — 
14, 22, same but nr Tarasovo, nr Gardeners’ partner- 
ship Zvezdnoye, 29.ix.2013. — Vitebsk: 14, Dokshitsy 
district, nr station Krulevshizna, 20.x.2000, pine forest 
(L. Chumakov). 


Bactericera ? femoralis (Foerster, 1848) 


Material examined: Vitebsk: 19, Dokshitsy dis- 
trict, station Krulevshizna, 20.x.2000, pine forest 
(L. Chumakov). 


Comment: Due to the poor state of the specimen at 
hand the identification is questionable and it could be 
also Bactericera acutipennis or B. bohemica. 


Bactericera reuteri (Sulc, 1913) 
Material examined: Gomel’: 24, Khoyniki district, 
Orevichi, Pripyatskiy National Park, 23.vii.1991, 
Malaise trap. 


Bactericera striola (Flor, 1861) 


Records: Recorded by Loginova (1962b) as Trioza stri- 
ola from the Vitebsk region. 


Bactericera substriola Ossiannilsson, 1992 


Material examined: Minsk: 14, Minsk district, 
Shchemyslitsy, Dubrava Natural Monument, 4.vii.2002, 
on Salix sp. — 14, same but Molodechno district, station 
Shipulichi, river Zapadnaya Berezina bank, 27.vi.2002. 


Trichochermes walkeri (Foerster, 1848) 
Records: Recorded from Belarus by Petrov (2004). 


Material examined: Minsk: 14, Molodechno district, 
nr Vyazynka, 29.vii.2001. 


Trioza anthrisci Burckhardt, 1986 


Material examined: Minsk: 14, 19, Minsk district, 
nr Tarasovo, nr Gardeners’ partnership Zvezdnoye, 
29.1x.2013, on conifers (L. Serbina). — 19, same but 
Molodechno district, station Shipulichi, canal bank, 
27.vi.2002. — Vitebsk: 164, Postavy district, lake 
Chetvert’ south bank, 10.vii.2005. 


Trioza apicalis Foerster, 1848 


Records: Recorded from Belarus by Sidlyarevich & 
Bolotnikova (1992). 


Trioza cerastii (Linnaeus, 1758) 


Material examined: Minsk: 19, Volozhin district, nr 
Kaldyki, 11.vii.2007, mixed forest (L. Serbina). 


Trioza flavipennis Foerster, 1848 


Material examined: Minsk: 19, 39, Minsk dis- 
trict, nr Aksakovshchina, Wildlife sanctuary Podsady, 
28.1x.2013, mixed forest, on conifers (L. Serbina). 


Trioza galii Foerster, 1848 


Material examined: No locality data, 14 (S. Buga). 


Trioza proxima Flor, 1861 


Material examined: Minsk: 19, Minsk district, 
nr Aksakovshchina, Wildlife sanctuary Podsady, 
28.1x.2013, mixed forest, on conifers (L. Serbina). 


Trioza remota Foerster, 1848 


Material examined: Gomel’: 49, Zhitkovichi district, 
Pripyatskiy National Park, 20.x.1998, pine forest 
(L. Chumakov). — Minsk: 24, 39, Minsk district, 
Shchemyslitsy, Dubrava Natural Monument, 26.1x.2013, 
on conifers (L. Serbina). 


Trioza urticae (Linnaeus, 1758) 


Material examined: Gomel’: 9%, 109, Zhitkovichi 
district, Krasnosel’ye, Pripyatskiy National Park, 
21.v.1991, Malaise trap. — 14, 69, same but Orevichi, 
21.v.-18.vi.1991. — 84, 209, same but 23.vii.1991. — 
Minsk: 34, 39, Logoysk district, Kupalovskiy National 
Park, nr Besyady, river Udra floodplain, 26.v1.2002, 
on various plants including Urtica dioica. — 14, 19, 
same but Minsk district, nr train station Kryzhovka, 
river Poplav bank, 3.vi.2001, floodplain meadow. — 


14 L. Serbina, D. Burckhardt & O. Borodin 


14, same but nr Chirovichi, hill, 9.vi.2002, dry mea- 
dow. — 134, 49, same but Shchemyslitsy, Dubrava 
Natural Monument, 15.v.2008, on Urtica dioica (L. 
Serbina). — 14, same but 15.viii.2008. — 1 adult without 
abdomen, same but 25.vi.2009. — 24, 29, same but 
Botanical Garden (Minsk), 16.vii.2009. — 24, 149, 
same but Shchemyslitsy, Dubrava Natural Monument, 
27.vii.2009. — 44, 29, same but 26.ix.2013, on conifers. 
— 14, 19, same but nr Aksakovshchina, Wildlife sanc- 
tuary Podsady, 28.ix.2013, mixed forest. — 14, same 
but nr Tarasovo, Gardeners’ partnership Zvezdnoye, 
29.1x.2013. — 74, 39, same but on conifers. — 24, 79, 
same but Molodechno district, nr Vyazynka, 16.vi.2001. 
— 14, 19, same but 17.vi.2001, alder forest. — 34, 49, 
same but nr train station Shipulichi, river Zapadnaya 
Berezina bank, 27.vi.2002. — 19, same but nr Sitsevichi, 
riverbank, behind station, 18.v.2007. — 32, same 
but river Udranka bank, nr bus stop, 2.vii.2008 (L. 
Serbina). — 14, 19, same but Myadel’ district, nr 
Naroch’, 4.vii.2009, on Urtica dioica. — 8%, 102, same 
but Volozhin district, nr Kaldyki, 11.vii.2007, mixed 
forest. — Vitebsk: 24, Dokshitsy district, Krulevshizna, 
20.x.2000, pine forest (L. Chumakov). 


Trioza velutina Foerster, 1848 


Material examined: Minsk: 14, Molodechno district, 
nr Sitsevichi, riverbank, behind station, 18.v.2007. 


DISCUSSION AND CONCLUSION 


Despite the fact that the psylloid fauna of Central and 
Eastern Europe is generally well studied (Gegechkori & 
Loginova, 1990; Klimaszewski, 1975), little is known 
from Belarus with only 12 recorded species one of which 
is doubtful, i.e. Aphalara polygoni (Loginova, 1961, 
1962b; Palyakova, 1969; Byazdzenka e? al., 1973; Gor- 
lenko ef al., 1988; Sidlyarevich & Bolotnikova, 1992; 
Petrov, 2004, 2011; Petrov et al., 2011; Petrov & Sautkin, 
2013). Based on recent collections we confirm the pre- 
sence of 4 previously recorded species and add 43 spe- 
cies, bringing the number of confirmed Belarusian psyllid 
species to 54. The majority of these are widely distributed 
in Europe and slightly more than half of them are asso- 
ciated with woody plants. The rest of the species is either 
introduced (Cacopsylla pulchella) or represents possible 
boreal elements (“peatbog” species as Cacopsylla ledi, 
Psylla betulae, Bactericera acutipennis). Craspedolepta 
crispati 1s rare and has been recorded from only a few 
countries (Lauterer & Burckhardt, 2004). 

The 54 species represent probably less than half of 
the number of species existing in the country by 
comparison with the surrounding countries: Poland 
112 spp. (Klimaszewski, 1975; Gtowacka, 1989, 1991; 
Glowacka & Migula, 1996; Drohojowska & Glowacka, 
2011; Kuznetsova ef al., 2012), Northwest Russia 47 


spp. (Loginova, 1954, 1961, 1962a, b, 1966, 1967, 1968, 
1972a, b), Lithuania 12 spp. (Vengeliauskaite, 1974; 
Malumphy ef al., 2009) and the former Livonia (now parts 
of Estonia and Latvia) 43 spp. (Flor, 1861) (Appendix 
2). There are no published data available for the psyllid 
fauna from the North of Ukraine and it is, therefore, not 
included in Appendix 2. Based on the occurrence of 
their respective host-plants following additional species 
(not listed in Appendix 2) can be also expected to occur 
in Belarus: Aphalara longicaudata Wagner & Franz, 
1961, A. polygoni Foerster, 1848, A. ulicis Foerster, 
1848, Craspedolepta campestrella Ossiannilsson, 1987, 
C. innoxia (Foerster, 1848), Arytaina maculata (Löw, 
1886), Cacopsylla affinis (Löw, 1880), C. breviantennata 
(Flor, 1861), C. picta (Foerster, 1848), C. rhamnicola 
(Scott, 1876), C. rhododendri (Puton, 1871), Psylla 
betulaenanae Ossiannilsson, 1970, Trioza scottii Löw, 
1880. 

To stimulate further research on Belarusian psyllids we 
provide here an illustrated identification key for 127 
species whose occurrence in the country has been doc- 
umented or is likely given that they have been reported 
from adjacent countries (Appendix 2) or their host-plants 
occur in Belarus. Targeted field work sampling on poten- 
tial host-plant is necessary to find also the more localised 
and rarer species. 


ACKNOWLEDGEMENTS 


We thank S. Buga, A. Egiyan, V. Karasev, E. Shesta- 
kov, L. Chumakov and Y. Gerashchenko for collecting 
part of the material discussed here. We are also grate- 
ful to L. Costeur, S. Buga, O. Nesterova and F. Sautkin 
for valuable advice to LS during her study. The critical 
comments and valuable suggestions on a previous manu- 
script draft by J. Hollier and I. Malenovsky are gratefully 
acknowledged. This study was partially funded by a grant 
of the Swiss Confederation (Federal Commission) to LS. 


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16 L. Serbina, D. Burckhardt & O. Borodin 


APPENDIX 1 


Identification key to adult psyllids of Belarus 
(Cacopsylla merita and Eryngiofaga deserta are not included in the key as their occurrence in Belarus is unlikely). 


l Vein R-M--Cu of forewing: bifurcating into veins RiandMtiCuUgi TTT eee 2 
Vein R+M+Cu of forewing trifurcating into veins R, M and Cu. — Triozidae (Fig. 2)... 84 
2 Metacoxa slender with flattened, tubercular meracanthus. On Acer — Aphalaridae, Rhinocolinae .......................... 
ORT Rhinocola aceris 
Metacoxamassıyeswithldistuinetgspur shapedimeracanthu se 3 
3 Vertex longer than wide. Segment 2 the longest antennal segment. On monocots (Carex, Juncus). — Liviidae, Livi- 
navi <A LELE GIO ERE 4 
_ Vertex shorter than wide. Segment 3 the longest antennal segment. On dicots ....................... i 6 
4 Vertex, in dorsal view, forming narrowly rounded anterior lobes with deep median cleft between lobes. Forewing 
short, oval, with evenly curved anterior margin, bearing indistinct dots in apical half. On Juncus........ Livia junci 


= Vertex, in dorsal view, forming broadly rounded anterior lobes with shallow indentation between lobes. Fore- 
wing longer, oblong oval or rhomboidal, in the middle with almost straight anterior margin, pattern different. On 
CON EX Sn dello ahi BR Ba ao e LI ann LOTO TO 5 
3 Forewing with subparallel fore and hind margins, without dark band along apical and anal margin. On Carex ..... 
LR ERI AE DIETA E RISI Ehe Livia crefeldensis 
— Forewing widening towards apical third, with broad dark band along apical and anal margin. On Carex .............. 
Se FOREST ARETE EEOC EEC EEOC MOTTA OI EEE Livia limbata 
6 Basal spine of metabia always absent; apical metatibial spurs spaced in equal intervals, forming open crown; if 
grouped, then vertex flattened and rectangular with anterior lobes, or evenly passing into genae ......................... 7 
~ Basal spine of metatibia often developed; apical metatibial spurs always grouped. Head with distinct, though 
sometimesismaliscenaliprocessestaesylitda e ME SY, A eee eee 37 
7 Head bearing genal processes. On Fraxinus — Liviidae, Euphyllurinae, Psyllopsis ........... iii 8 
FICadiwithoutKecnaliprocessesSoenAeIrOUNdedFanteri OI 11 
8 Body green, forewing lacking dark pattern. Paramere axe-shaped (Fig. 3). Female proctiger distal to circumanal 
ring abruptly tapered (Fig: 4). RS FERN Psyllopsis fraxinicola 
— Body with dark areas, forewing with dark veins and more or less expanded dark pattern. Male and female termi- 
nalia different... GIL an ani LI RS SERIE NER nn 9 
9 Forewing lacking continuous marginal band apically; membrane dark at apices of veins M and Cu, in the middle 
of vein Cu,, and along vein Cu,, (Fig. 5). Thorax yellow-orange, dark brown dorsally. Paramere, in profile, ham- 
mer-shaped with large anteriorly directed lobe (Fig. 6). Apex of female proctiger broad (Fig. 7). .......................... 
BILE PRIORI TENCE OCT RONDE EEE ee Psyllopsis discrepans 
— Forewing with dark band along apical margin stretching from fore to hind margin (Figs 8, 11). Thorax yellow with 
brown or black pattern. Paramere, in profile, with posteriorly directed lobe (Figs 9, 12). Apex of female proctiger 


narrow.:(Figs 10” 13)= eee A SR Te TTT 10 
10 Forewing pattern very dark, extended and well-delimited (Fig. 8). Paramere with large rectangular posterior lobe 
(Fig. 9). Female proctiger with slight hump distal to circumanal ring (Fig. 10)................. Psyllopsis distinguenda 


— Forewing pattern slightly lighter, more reduced, often forming only narrow band, and less clearly delimited towards 
interior (Fig. 11). Paramere with small triangular posterior lobe (Fig. 12). Female proctiger straight or concave 


distalitocircumanalrine (Riel) PR Re RSR Psyllopsis fraxini 
11 Metabasitarsus without black spurs. On Populus — Liviidae, Liviinae ............................. Camarotoscena speciosa 
— Metabasitarstis with'2?blackspürsteer er ee E OR EEE 12 
12 Male proctiger without posterior lobes (Fig. 14). ALHW always < 1.0. On Calluna vulgaris. Liviidae, Euphyllu- 
Saree ee eee RO Te Strophingia ericae ' 
— Male proctiger with long, wing-like posterior lobes. ALHW often > 1.0. — Aphalaridae, Aphalarinae................ 13 
13 Vertex with distinct angular anterior lobes which are separated by narrow transverse groove from genae. Clypeus 
more or less distinctly protruding from lower head surface. — Aphalara............................................................. 14 
_ Vertex with weakly developed anterior lobes, smoothly passing into genae. Clypeus short, pressed against lower 
headisunfaceSnotistrongly4protrudingstromienael@rasp 40 DA 24 
14 Forewing with dark pattern consisting of well-defined spots or patches. Apical dilatation of aedeagus with large 
dorso-apical membranous sack. Circumanal ring of female proctiger never expanded caudally ......................... 15 


— Forewing without well-defined dark pattern but sometimes infuscate. Apical dilatation of aedeagus without or with 
small dorso-apical membranous sack. Circumanal ring on female proctiger usually expanded caudally (exceptions 
A. longicaudatas:A .\purpurGS CONS) eis. fee 20 RE OI LEO 17 


15 


DI 


22 


23 


24 


25 


26 


27) 


28 


29 


30 


31 


32 


Jumping plant-lice of Belarus 17 


Clypeus long, clearly visible from above; cylindrical, constricted subapically. Forewing narrow, with dark patches 


concentrated in apical third, basal half clear. On Polygonum ................ iii Aphalara maculipennis 
Clypeus short, not or hardly visible from above; conical, apically blunt or subacute. Forewing broad, with dark 
DatchesfextendinstalsopntoJbasallH al EHE RE nn satesbeeue ic cialis toate eseen: 16 
Surface spinules of forewing forming short rows of 2-4 spinules (Fig. 15). Paramere produced apico-posteriorly 
(TOM (OT RU MER ANITA POLAT TROIA POT ICT Aphalara exilis 
Surface spinules of forewing forming longer rows (Fig. 16). Paramere not produced apico-posteriorly (Fig. 18). On 
ULI CN VAIANO LE ORO TOTI Aphalara ulicis 
Head and thorax dark brown or black. On Sfellaria graminea ..................-. ii Aphalara affinis 
Kleadtandithoraxgochreousswithloran gelonbro wis Han Se 18 
Forewing with surface spinules arranged in irregular, transverse rows (Figs 19, 22-23)... 19 
Surface spinules of forewing arranged in irregular squares or rhombi (Figs 24-26) ........... i 22 
Paramere with posterior extension apically (Fig. 20). Female terminalia long (Fig. 21). On Polygonum bistorta .. 
ee ODA OOE ERA PEER EEURP ERA ER PE RP PEN CPE SPP EE PP EE PEER EEE IT OU PEU PESTE ES Aphalara longicaudata 
Barameremotiextentedipostero-apicallARemaletemiNANAISNOTE ER RESSE E 20 
Body dimensions large (length from head to apex of forewing when folded over body > 2.9 mm). Forewing mem- 
brane amber-coloured, surface spinules arranged in very dense, transverse rows (Fig. 22). On Caltha .................. 
PARATA AE TRI EEE EU SE REESE RPE UE UT EU Ee Aphalara calthae 
Body dimensions small (< 3.2 mm). Forewing membrane colourless or fumose, surface spinules arranged in sparse 
TOWS]((Ri 0928) PR Re CR RME find ee eee 21 
Circumanal ring caudally consisting of several rows of pores. On Polygonum, Rumex............ Aphalara polygoni 
Circumanal ring caudally consisting of two rows of pores. On Rumex ............. Aphalara purpurascens 
Male paramere with anterior finger-like process situated subapically (Fig. 27). Aedeagus as in Fig. 28. Female 
(TTT, GS WH IE, DS), Oi LEO Lo III RIA II IO IE SROO, Aphalara borealis 
Male paramere with anterior finger-like process situated close to apex (Figs 30, 33). Aedeagus as in Figs 31, 34. 
Rene (STRATOS TT JONES ES SR I MR Ti rt nn nn eee 23 
Tip of distal portion of aedeagus directed in an angle of about 30° to longitudinal axis of segment (Fig. 31). On 
ROIS ONUIMIGVICILALCLOTOUP emcee eet TR te site reset Aphalara avicularis 
Tip of distal portion of aedeagus directed in an angle of about 90° to longitudinal axis of segment (Fig. 34). On 
ROL ONU MERE ER MIR Re SI IRE EEE EEE Aphalara freji 
Forewing with pattern consisting of well-defined, dark spots of 10-30 um diameter... DS 
Forewing without pattern, or with pattern consisting of dark bands, or membrane irregularly infuscate ............ 29 
Body bearing macroscopic setae which often are covered in wax and thus resemble scales. On Artemisia abrota- 
LOI. ROOT OAV I Craspedolepta alevtinae 
B OdYA LAC KIN PIN ACT OSCO PICIS ELA E ER IAT II IRR 26 
Forewing with surface spinules completely covering membrane (Fig. 36). Terminalia as in Figs 41-42. On Arte- 
VILLS Docc be cocc ER CEE ESE Re Re An DESSERO PERCE Craspedolepta artemisiae 
Forewing with surfaces spinules forming hexagonal pattern (Figs 37-38)... DY 
Forewing with dark spots densely spaced, partly confluent in apical part. On Artemisia campestris «2.1.1.1 
Beet ECE EEOC SERRE ee eee CEE ee eens eee Craspedolepta campestrella 
Forewing with dark spots sparsely spaced, not confluent in apical part of wing (Figs 39-40)... 28 
Dark spots on forewing dark brown or almost black; surface spinules relatively sparsely spaced (Fig. 37). Termi- 
NAM AASMINPRI SSIS 44M ONVALICMNISICMVU CAG SIRO RI RI IRR RAI RO Craspedolepta latior 
Dark spots on forewing pale to yellow brown; surface spinules relatively dense (Fig. 38). Terminalia as in Figs 
AS — Om OMPALLCIIISLALADSINLNUIM GA ame Craspedolepta malachitica 
Male paramere with large apical triangular posterior lobe, and subapical anterior process which is very long and 
directed backwards (Fig. 47). Female proctiger ending in two points (Fig. 48). On Chamerion .......... ui 
96095900005 FIORITO NINNI IRRITANTI TIT II Eee eee eee eee eee Craspedolepta subpunctata 
Male paramere, in profile, club-shaped, or if triangular, then anterior process not directed backwards (Figs 50, 53, 
SU Ol, ©3, CD=GO)) ETNA PIOCUILEMENTINLANIAIS IN (9 [00110 I 050535050 neretto 30 
Body length from head to apex of forewing when folded over body < 3.2 mm ....... 31 
B'OUVEEN LINE RSI ME INI IO COLE EEE ACEO EE EER GREE EE EPEC eee Eee EEE RR 35 
HOC Win Paw thOUtdarke pal ChestOrMStrl PESTE Un rennes nee 52 
am PANTIN El IOMATOIDIONMIPALCRESIONS TAPER. 58 


Surface spinules of forewing arranged in irregular transverse rows (Fig. 49). Male paramere with large triangular 
apex and long straight anterior process on inner surface (Fig. 50). Female proctiger more than 4 times longer than 
po rms lenzii (a SI) KOT CS ee eee Craspedolepta innoxia 


18 


36 


L. Serbina, D. Burckhardt & O. Borodin 


Surface spinules of forewing widely spaced, sometimes partially reduced not forming transverse rows (Fig. 52). 
Male paramers club-shaped, with short, claw-like anterior process (Fig. 53). Female proctiger less than 4 times as 
long as pore ring length (Fig. 54). On Artemisia vulgaris .…....................................,........... Craspedolepta omissa 
Forewing pattern with dark brown, well-delimited patches forming bands along outer margin and in the middle 
stretching between the apices of veins R, and Cu,,, and a spot in cell cu, (Fig. 55). Male proctiger with broad wing- 
like posterior processes lacking a basal hook (Fig. 56). Paramere as in Fig. 57. Female subgenital plate suddenly 
Natrowedisubapicallya (Riess) 5 Oni Cam eri Or RR Craspedolepta nebulosa 
Forewing pattern ochreous to brown, forming bands along the veins in apical half or a band along wing margin. 
Male proctiger with narrow single-like processes bearing a basal hook. Paramere different, with rounded to angu- 
lanapicalidilatationskemalesubeenitaliplatetevemlyatapenecl EEE 34 
Antenna usually 8-segmented. Forewing pattern distinct, restricted to narrow stripes along veins in apical half 
(gup 9S9) ierminaliafasiniBigs(OC02A0NH CARRA Craspedolepta nervosa 
Antenna usually 10-segmented. Forewing pattern forming a band along outer wing margin (Fig. 60). Terminalia as 
IniFipsi63-6430OnAchillegt St: caio MIRATO SOIN RO E EROE ne Craspedolepta bulgarica 
Anterior margin of vertex strongly indented in the middle with two distinct tubercles anteriorly; antero-lateral 
margin of vertex dorsal of antennal insertion distinctly concave. Forewing yellowish to brownish ochreous. On 
ECONOdON: irs OTA OEE Craspedolepta flavipennis 
Anterior margin of vertex weakly indented in the middle with two indistinct tubercles anteriorly; antero-lateral 
margin of vertex dorsal of antennal insertion more or less straight. Forewing semitransparent to whitish.......... 36 
Forewing veins dark brown, membrane light, sometimes with light brownish spots or brownish tinge apically. 
Paramere stalk robust, apical spoon-like part shorter and rounded as in Fig. 65. Female terminalia as in Fig. 67. On 
Seneciorintegrifollus: <a ra EE Craspedolepta crispati 
Forewing veins concolorous with membrane, membrane with yellowish tinge and brownish dots in apical third. 
Paramere stalk slender, apical spoon-like part longer and subangular as in Fig. 66. Female terminalia as in Fig. 68. 


Oni LeGniodon..s rss RS O E O Craspedolepta sonchi 
Metabasitarsus with only one outer black spur, rarely also with much smaller inner spur. Propleurites divided by 
Vertical: SUture:.;.;. ata ia ee E O I TET 38 
Metabasitarsus with two well-developed black spurs which are subequal in size. Propleurites divided by diagonal 
SUfUTE siii n e EES 44 
Either genal processes longer than vertex, or length ratio of veins Cu / Cu, of forewing 0.9-1.1. Paramere usually 
slender.— Livilla rise RS I TO IE SERRE 2. 39) 
Character combinationdifferent::.2......:...n2.22 fe Re TO RER Re De 41 
Rorewing ovalsstrongly;convexsconaccouss On Gyiisusy GCHISTA PIRATI ARIA IATA Livilla ulicis 
Rorewinstoblong{ovalStatsmembranousi(bIs00 7) ARI 40 


Forewing long and narrow, branches of vein M forming an acute angle, pattern as in Fig. 69. On Genista ........... 
nia RI ORIO RR e °° Livilla horvathi 
Forewing short and broad, branches of vein M forming a right angle, pattern as in Fig. 70. On Chamaecytisus, 


CIISUS: sii ete AR PRE M RE ORNL NOT ETE RE Livilla radiata 
Forewing broadest in apical third, lacking colour pattern. Paramere very slender, curved caudad, digitiform api- 
cally. Dorsal margin of female proctiger, in profile, concave. On Cyrisus scoparius ......... Arytainilla spartiophila 


Forewing broadest in the middle or in basal third, with conspicuous brown pattern. Paramere broader, paral- 
lel-sided, in profile, truncate apically with large, forward pointing apical tooth. Dorsal margin of female proctiger, 


In profile; Straight. — Arytalna es RR ee ETS 42 
Forewing broadest in the middle, fore margin relatively flat. Paramere broad, dorsal margin distinctly concave. On 
Cytisus; Genistiz:: cc ARRE IE a Arytaina genistae 
Forewing broadest in basal third, fore margin strongly curved. Paramere narrow, dorsal margin weakly convex. On 
Chamaecytisus:ratisbOnensis RSR RE EN IR O. Arytaina maculata 
Antennal ségment9 longer, than’ 10:58 Re co 44 
Antennal segment, 9:shorterithan l'O. 2 EN een 48 
Genal processes short and broad. Forewing with costal break and pterostigma developed; membrane yellow, veins 
concolorous. Dorsal margin of female proctiger serrate in apical third. On Alnus .................. Baeopelma foersteri 
Charactencombinationyditikerem tae 27251 cement Rene nee 45 
Forewing with cell cu, almost as high as long; membrane yellowish, veins concolorous, yellow or green. On 
BUXUS ur OE TL A ES CRT Psylla buxi 
Forewing with cell cu, at most 1.5 times as long as high. On Betulaceae .......................... i 46 
Horewinetlackinecostalibreakfandipter ose MAO AUS Psylla fusca 


Korewinsibeaningscostallbreakgandipterosti In A aera eee ee 47 


47 


48 


49 


50 


55 


56 


57 


58 


59 


60 


61 


62 


63 


Jumping plant-lice of Belarus 19 


Genal processes short, rounded. Forewing with dark brown or black veins in apical two thirds. On Alnus ............ 
5000003003000 S CCAS HOCGE OCC BEOSEEC IAA FIATI PILE AI AIAR peer Psylla alni 
GenaliprocessesilongermconicalahorewinPavcinsipaler ONE 48 
Body length, including forewings when folded over the body, longer than 3.5 mm in males and 4.1 mm in females. 
Rorewin cco ounlesstormuathby.clLowavin LEONE CIO seen SNO nearer e Psylla betulae 
Body length, including forewings when folded over the body, shorter than 3.7 mm in males and females. Forewing 
AMDE COLE ONE CH GRR an en Rie Psylla betulaenanae 
Metatibiaswithl4yapicalisp Urs NON PB CUIR Chamaepsylla hartigii 
Metatibiakwathtsrapicalispurs ACTOR Recon eee ni 50 


Dorsal surface spinules in cell r, of forewing above bifurcation of vein M arranged in squares or rhombi of about 
20 um length; surface spinules in cell c+sc restricted to apical portion of cell or entirely reduced; surface spinules 
reduced in basal part of r,, at most a few spinules present; fields of surface spinules tapering along apical wing 
margin; forewing membrane always colourless; pterostigma oblong cuneate, evenly tapering. Antenna shorter than 
1.75 mm, if longer, then forewing longer than 3 mm. Male paramere simple, lamellar. Female terminalia short, 
(CMOS onn00 20000000 DOCS RER RE OI IIS 51 

Forewing spinulation different, or wing membrane yellowish or brownish, or pterostigma elongate with subparal- 
lel margins. If antenna longer than 1.75 mm, then forewing shorter than 3.0 mm. Male paramere often complex. 


Kemalestenmina MAFIE TI A ne Re Tha IRE 54 
ntennaishortenmthanl2imMiON CRALCAZIS ccoccccccccececooccxcc0e00c059000000000000000000000000000009000000000005 Cacopsylla peregrina 
Atenas On Gergth anwl espn MPEREZ RE RI E O TT TRE SL 
Antenna oneergtha I WSIMOIONU IL RI RI SR RIO I Cacopsylla ulmi 
Ain tennats None NANTES SENTE ER RI OI RT I O ON eee Ra 58 
Antennalsegmentsg4-Sswithiblackgapex ONE 0 D USER nee Cacopsylla sorbi 
Antennal segments 4-8 with yellow or ochreous apex. On Malus Us Cacopsylla mali 
Dorsal surface spinules of forewing in cell r, above bifurcation of vein M irregularly, densely spaced (2-10 um) or 
ATEATI OCC RITINEATIS V CTSCITOW Stee Semen TOTO an nl le er cans 55 
Dorsal surface spinules of forewing forming more or less even squares or rhombi of 20 um distance in cell r, above 
DILULCALONIO MVC E RR nine ROCHE RE EERE CECE eee eee 62 
SUMS SOMOS GIR PEA IAN SVEN ENI ON S.ccc0q000cc0G000c00000000000000000000000000060000000090000000000000.000000000009000000000000000000000000 56 
Sustacesspinulesidenselysinze LULATIVES DAC EC ISAIA se 58 
Paramere, in profile, with large basal lobe. Dorsal margin of female proctiger straight or weakly convex. On 
NADA RIN RITI D ESET iI ee reni cala re eu Luton Cacopsylla elegantula 
Paramere, in profile, lamellar with anteriorly directed apical tooth. Dorsal margin of female proctiger sinuous. On 
IS III ISO III ITER TTT 57 
Thorax brown, abdomen green. Terminalia ochreous or brown. On Salix............ Cacopsylla abdominalis 
BodYColounevEn VAS hHOMEATSAAONIS A LEE Cacopsylla intermedia 
Forewing with an apical dark brown band or completely brown to dark brown... 59 
Rorewin ele ia amos Ve llO WIOAOCNTE O IS PORT oe 60 


Forewing bearing dark band apically (Fig. 71). Metatibia without genual spine. Male subgenital plate bearing api- 
cal tubercular extension (Fig. 72). Female proctiger and subgenital plate ending in thorn-like processes (Fig. 75). 
OES O7 DS eee IIa ani Cacopsylla breviantennata 
Forewing irregulary dark without distinct apical ribbon (Fig. 73). Metatibia with genual spine. Male subgenital 
plate rounded apically (Fig. 74). Female proctiger and subgenital plate evenly tapering in profile (Fig. 76). On 


IR HUIT SERRE RE RO RR DIE n EEE EEE PER COC EEE ei n LL LA aa Cacopsylla pruni 
Forewing bearing ventral surface spinules in cell c+sc. Surface spinules on forewing very densely arranged. On 
IS CI LG ara cla SR ori el CECE CEC CECE EE le lalui Cacopsylla ambigua 
Ventral surface spinules lacking in cell c+sc. Surface spinules on forewing more sparsely arranged.................. 61 
Forewing oval, widest in the middle; wing apex near apex of vein M,.. Antenna shorter than 1.0 mm. On Salix 
REES ES MOS MANN LLC meron ee Oe din ARES Cacopsylla parvipennis 
Forewing widest in apical third; wing apex at the middle of outer margin of cell r,. Antenna longer than 1.0 mm. On 
SALARIA I are i arco Cacopsylla flori 
Forewing bearing dark brown patches on tips of veins, at the bifurcation of vein M, and in the middle of vein Cu, .. 
OMOCLCISIS GUASTI NE EEE Cacopsylla pulchella 
Boreywinsspattermgdihienen EME IR RE RI O NI TE 63 


Dorsal surface spinules covering entire cell c+sc of forewing apart from stripes along veins; forming extended 
fields in other cells which taper towards wing margin; membrane colourless or fumate but never with brown, 
THON COMMAS Saye ALOT WEL U ARE ER 64 


64 


65 


66 


67 


68 


69 


70 


L. Serbina, D. Burckhardt & O. Borodin 


Dorsal surface spinules of forewing more or less reduced, or not tapering towards wing margin, or wing pattern 
consistingswithidarky strong lyicontrastedistmpetal on gave CURE ER 67 
Antenna usually longer than 1.2 mm. Genal processes broad and blunt. Paramere broad, lanceolate (Fig. 77). Dor- 
sal margin of female proctiger, in profile, weakly sinuous, apex narrowly rounded (Fig. 78). On Pyrus ................ 
silla NIRO ARRE SR SL,  ——————t—t—m€—m Cacopsylla pyrisuga 
Antenna usually shorter than 1.1 mm. Paramere narrow or with apical processes. Dorsal margin of female procti- 
gericoneayelinitheimiddleTorzapexsttun EAT E Re 65 
Paramere, in profile, with square base bearing 2 apical processes (Fig. 79). Dorsal margin of female proctiger, 
distal of cireumanal ring, evenly concave; apex rounded (Fig. 80). On Malus........... iii Cacopsylla picta 
Paramere, in profile, elongate; apex with inward and forward pointing tooth. Dorsal margin of female proctiger 
raised'inthe middle apexcan oul ar: 2258 ‘io 6ee gt ttt, o 01 66 
Paramere long, in profile, narrowed in the middle. Apex of distal segment of aedeagus weakly curved (Fig. 81). On 
CROAEGUS ZE ESRI, RIO SII Cacopsylla melanoneura 
Paramere short, in profile, evenly tapering from base to apex. Apex of distal segment of aedeagus strongly curved, 
hook-shaped'(Fig382)- Oni Crataegus). 0 RR O O Cacopsylla affinis 
Forewing with fields of surface spinules tapered along apical wing margin; membrane bearing dark brown patch 
alon EC (Res ER) Soon RESSE PROSE SENSORE SIRIA EEE n. 68 
Character combinationidifferenti.:-= sala il nn Ronn ———_—c 69 
Areas of radular spinules of cells m,, m, and cu, of forewing light; dark patch along vein Cu,, not reaching bifur- 
cation of Cu, not narrowed in Drome half (Fiz. 83); surface spinules forming extended fa: in cells ctsc andr.. 
OS OF DUS ER RSR OE >> Cacopsylla ns 
Areas of radular spinules of cells m,, m, and cu, of forewing more or less dark; dark patch along vein Cu,, reaching 
bifurcation of Cu, straight in proximal half (Fie. 84); surface spinules reduced in cells ctsc and r,. On Gr ataegus. 


RAT oa suas ZAR RIO RO C ete crataegi 
Surface spinules of forewing forming very narrow fields. On Rhamnus ........... Cacopsylla rhamnicola 
SurfacexspinulestofforewinesfoL min szextend edle |d SORA 70 
Forewing with fields of surface spinules tapering towards apical margin; clavus with brown apex. ................... 71 
Character.combination différent: RIN, RT Rs 12 
Paramere sickle-shaped (Fig. 85). Female proctiger strongly narrowed in the middle (Fig. 86). On Pyrus............. 
GNA ARR ORI AI O —— ——_ Cacopsylla pyri 
Paramere lamellar (Fig. 87). Female proctiger cuneate (Fig. 88). On Pyrus .......... i Cacopsylla pyricola 
Anténnatlongerithantl:6 mms ee 73 
Antenna:shorter:than:1:3:mmici er TRS A RE ee SR AR Cc cae 15 
Metatibia with 1+1+(2-3)+1 sclerotised apical spurs. Antennal segments 3-7 yellowish or ochreous with dark 
brown apex. Fields of surface spinules tapering along apical wing margin. On Viscum ................ Cacopsylla visci 
Character combination different... TTT n. 74 
Body colour dark brown. Paramere with short, angular apical, sclerotised apex (Fig. 89). Valvula 2 of female ter- 
minalia with straight ventral margin (Fig. 91). On Hippophae …........................................ Cacopsylla zetterstedti 
Body colour green or yellow. Paramere with long, curved apical, sclerotised apex (Fig. 90). Valvula 2 of female 
terminalia with concave ventral margin (Fig. 92) On Hippophae..…..................................… Cacopsylla hippophaes 
Pterostigma cuneate, broad and short, with converging margins ending in the middle of vein Rs; wing membrane 
vellowishioRfochreoussveinsfochreousionlichtWDTo v1 76 
Pterostigma long and narrow, with subparallel margins ending in apical third of vein Rs; wing membrane colour- 
less:or:dark, veins light'ordark:....—« l'e re titti i MN CEE 79 
Fore margin of forewing relatively straight. On Rhododendron. ...........iiiii Cacopsylla rhododendri 
Koresmarginlo MFOLEWINLISTTONLYAGUEVE FR V7) 
Surface spinules entirely covering cell c+sc of forewing. On Vaccinium ............uiii Cacopsylla myrtilli 
Surfacefspinulestabsentifromibasalithird'o ce 1lIG:SCIO MOT will LEE 78 
On.Vacciniummyrtillisc: di Re OO Cacopsylla fraudatrix 
On Ledum palustvé x: BSc LIRE ES III Cacopsylla ledi 
MaleiparamerelbearineSubapicalobelalon thin dima INTER 80 
Malejparamerellackingysubapicalllobelalon syhindam ATEN 82 
Raramerejlackingsbasalllobelawhindimarcins ON SA ARR Cacopsylla moscovita 
Raramere) bearingibasalylobeya tian bin Arg in rm 81 
Basal lobe at hind margin of paramere not incised dorsally. On Salix. ........ ii Cacopsylla saliceti 
Basal lobe at hind margin of paramere strongly incised dorsally. On Salix............ Cacopsylla iteophila 


Apex of paramere forming simple, backwards directed sclerotised tooth. On Salix ................ Cacopsylla pulchra 


83 


84 


85 


86 


87 


88 


89 


90 


91 


92 


93 


94 


95 


96 


Jumping plant-lice of Belarus 21 


ANDO Oi PALA EILEAVILMIEWOISITONLIVESCIETOLSE NEC RIENTRI RO NES TRTOre eretto 83 
Paramere, in rear view, with a tooth in apical third; in profile, as in Fig. 93. On Saliîx.......... ii 
Rens rade su denr ones eee isans ste are sasesessee Cacopsylla brunneipennis 
Paramere, in rear view, with lobe in the middle of inner margin; in profile, as in Fig. 94. On Salix 
Ne NE DANA AAT III DITE I TTT Cacopsylla nigrita 
NTefati D ai th ES ISC ICT OISE AA DICA LIS PUERTA RIAPERTO TT nno 85 
Metatibiaawitha2iSCIerotiSe A PICA IS DUT ennemie senti ele resettare 107 
Forewing with extended dark pattern consisting of small points; fore margin straight to concave between apices of 
veins Rs and M,., (Fig. 95). On Rhamnus cafhartica.….….................…....... Trichochermes walkeri 
Horewineapatternmiditierenthore marin CON EX SE TIOZ AIDER ne 86 
Forewing short and broad, angular apically; branching of vein M distinctly distal to Rs—Cu,, line. Antennal colour 
sttonalygeontrastedgsesment@yli shtagse MEN SAUT AR ere 87 
Forewing elongate, sometimes rounded apically; Rs-Cu,, line proximal to or on branching of vein M. Antennal 
COLOUTAV AT a | ER Snr RM nn neue as essen mini nn Suess costlosadesnuebeilyegiedsscteasevedesdaweeses 89 


Genal processes short. Paramere, in profile, strongly narrowed in apical quarter, forming anterior tubercle and 
posterior process (Fig. 96). Female subgenital plate truncate apically, consequently strongly convex dorsally (Fig. 
97). Terminal setae on antennal segment 10 subequal. On Valerianella.................................... Trioza centranthi 
Genal processes long. Male paramere in apical quarter regularly narrowing to apex (Figs 98, 100). Female subgeni- 
tal plate evenly tapering to apex, dorsal margin almost straight (Figs 99, 101). Terminal setae on antennal segment 
IOIStTONEIVAUNE UA RE E doa d0655000¢050 C00 naa cacca1C ad ge0 00 SoC Seer cee eee eee Eee Core 88 
Forewing without surface spinules. Male paramere robust, angular apically; distal portion of aedeagus with large 
apical hook (Fig. 98); female subgenital plate, in profile, with obtuse apex (Fig. 99). On Galium and other Rubia- 


CCA III PO COC CEA DEEE LE SERS rec eee EEE eee eee eee Trioza galii 
Forewing with surface spinules. Male paramere slender, curved apically (Fig. 100); female subgenital plate, in 
pronleswithrsmallisharpioothtatapexd (RE MON LOL U EEE Trioza velutina 
Forewing with surface spinules present at most in cell cu,. Male paramere long and slender, lamellar (Fig. 102). 
Female terminalia long, dorsal margin of proctiger more or less straight (Fig. 103). On Urtica ...... Trioza urticae 
Forewing with surface spinules present in all cells, covering smaller or larger areas... 90 


Terminal antennal setae strongly differing in length, both distinctly shorter than antennal segment 10; shorter seta 
very short, stout and truncate apically. Axes of genal processes diverging forward; genal processes relatively mas- 


sive and blunt, evenly narrowed. Body orange to reddish, sometimes brownish to black...............................0.... 91 
Terminal antennal setae different; axes of genal processes parallel, or genal processes slender and pointed or fusi- 
LOTMAOMDOUNAVELIOWIS NOMETEEN SN Re tt 94 


Surface spinules of forewing spaced at 5-10 um intervals or forming transverse rows in the middle of cell r, at level 
of branching of vein M, never reduced along outer wing margin. Male paramere basally robust (Fig. 104), female 


terminalia short with proctiger, in profile, dorsally straight (Fig. 105). On Hieracium ................... Trioza proxima 
Surface spinules of forewing forming regular rectangles or rhombes of 15-10 um distance, sometimes reduced 
alon sfoulerswinsamareın TRTTTANA dI TTI coccc0c0s0000000000000000000007800200000d0000000000 3605 3e0NDpCoDNEDTEGTNEGaNSoBNBScDAESTECETIGRCOS 92 
Male paramere with weak subapical constriction. Female proctiger cuneate, regularly tapering to apex, short, about 
IGS as lone as Cimounmernell warns, Gi MEZ cccc00200000000000000000000000000000000000000000000000000000000000000000550550500 Trioza foersteri 
Male paramere with strong subapical constriction (Fig. 106, 108). Female terminalia different ......................... 93 
Male paramere about as long as or longer than proctiger, apical portion bent inwards and weakly backwards (Fig. 
106). FPHW less than 0.8, proctiger truncate apically (Fig. 107). On Taraxacum ............u ii Trioza dispar 
Male paramere shorter than proctiger or apical portion straight, directed upwards (Fig. 108). FPHW more than 0.8, 
proctiger regularly narrowed apically (Fig. 109). On Hieracium.............................................. Trioza tatrensis 
Surface spinules of forewing in cell r, at level of branching of vein M densely spaced in a distance of 6-12 um... 
eee SIC SEC ee I TI nn sienne aie dessus is 95 
Surface spinules of forewing in cell r, at level of branching of vein M arranged in regular rectangles or rhombi in 
a distance of 10-20 pm ...................... WE e 97 


Forewing distinctly angular apically; vein Cu much more than twice as long as Cu,,, cell cu, flat. Male paramere 
slender in apical half; apex turned backwards. Female proctiger blunt apically. On Aegopodium........................... 

50500000020 300G0 000000 TOO IO erence TRO Trioza flavipennis 
Forewing irregularly rounded apically; vein Cu at most slightly more than twice as long as Cu „, cell cu, high. Male 
paramere massive in apical half; apex turned inwards or forwards (Fig. 110). Female proctiger pointed apically 
(RIO) RA RN RE eae CE LE eee ee TUE 96 
Male parameres with large antero-basal lobe. Distal aedeagal segment with large apical hook. Female subgenital 
plate short, truncate apically. Lateral abdominal setae present on tergites 3 and 4 in males, and 4 and 5 in females. 


(NO) 
N 


97 


108 


109 


L. Serbina, D. Burckhardt & O. Borodin 


Relative lengths of terminal antennal segment : shorter terminal antennal seta : longer terminal antennal seta = 1.0 
0:4:::0.7.. On Cardam ines Ste llanid ">, ARI TT Trioza rotundata 

Male parameres without distinct antero-basal lobe. Distal aedeagal segment with short hook. Female subgenital 
plate long, pointed apically. Lateral abdominal setae present on tergites 3 in males, 4 in females. Relative lengths 
of terminal antennal segment : shorter terminal antennal seta : longer terminal antennal seta = 1.0 : 0.3 : 0.8. On 
Valerianella tripteriQis, Re tre Trioza tripteridis 

Forewing with very long, sinuous vein Rs; membrane transparent, colourless, wing margin with dark dots at inter- 
sections of veins and near radular spinules. One terminal antennal seta less than three times shorter than the other 


seta. On RhGMAUS SRE errant I ESSE EI FRS Trioza rhamni 
Combination. of Characters différentes RE RE TR 98 
BodYalmosticomplele  idark ADO WNIOND ICE aaa an 99 
Body green, yellow or reddish; sometimes thorax dark but abdomen green ...................... nennen 100 
Forewing membrane dirty whitish. Apex of paramere pointing forwards. Female proctiger truncate apically. On 
SORT AR es VORREI RI roe ER 050040000090700000000000 Trioza saxifragae 
Forewing membrane ochreous. Apex of paramere pointing backwards. Female proctiger subacute apically. On 
ASI GNU ING] OF SENIOR III en De 0. Trioza schrankii 
Body straw-coloured or ochreous. Paramere simple, broadly lamellar. Female proctiger short, about twice as long 
AS CITEUMANALITIN LA ONERU M EXIS CUL ALU SERA TTT Trioza rumicis 
Body green, light yellow, sometimes thorax brown. Paramere with inner process or with broad base and narrow 
apical portion. Female proctiger about three times as long as circumanal ring .................\ 101 
Forewing broadly irregularly rounded apically. Male proctiger with posterior lobe; paramere with inner process 
(Fig... 112)..On.Gerastium:=..- RESERO RR Trioza cerastii 
Forewing angular apically. Male proctiger without posterior lobe; paramere without inner process................. 102 
Body always, also in overwintered specimens, yellow or green, never yellowish orange, or with brown to black 
SPOUS ws. svesvsceeescecsteacagecoucsouseespesnstencntedeveants che sui es teen I O ER 103 
Body never exclusively yellow or green; either yellow-orange or with brown to black spots ........................... 105 
Antennal segments 6-8 brown. Basal portion of male paramere with distinct antero-apical tubercle (Fig. 113). 
Female proctiger, in profile, with weakly sinuous dorsal margin (Fig. 114). On Cirsium .................... Trioza cirsii 
Antennallsepments{6-8ilishtMale andifemalelterminalialdiife enter 104 


Basal portion of male paramere angled antero-apically. Female terminalia long. On Cirsium......Trioza agrophila 
Basal portion of male paramere oblique antero-apically. Female terminalia short. On Senecio... Trioza senecionis 
Postero-apical process of paramere very slender and straight. Female subgenital plate much shorter than proctiger. 
ON CHIPS ANN CMU ie RIO OT Trioza chrysanthemi 
Postero-apical process of paramere wider and curved. Female subgenital plate about as long as proctiger...... 106 
Paramere with very small anterior lobe (Fig. 115). Female subgenital plate slender apically. On Achillea............. 

RE A AO II IA DIRI Trioza abdominalis 


Paramere with large anterior lobe. Female subgenital plate massive apically. On Knautia............... Trioza munda 
Branching of vein M of forewing distinctly distal to Rs—Cu,, line; vein Rs straight or concavely curved to fore 
margin; forewing angular apicallyerenereesseeesseesserseee ner eene see nn n 108 
Branching of vein M of forewing proximal to or about on Rs—Cu,, line, or vein Rs sinuate, or apex of forewing 
broadly'rounded...:.......r02.20002220 00er sets M ees eee eee ee RETO TE TSS ECOG n n. 110 
Genal processes short. Forewing very narrow with low cell cu,; surface spinules absent. Male paramere narrow 
(Bio) SremalerterminaltayassınaBi SAVA OWE US peeeeeneeeeeseeeeee eee nn Trioza alacris 
Genal processes longer. Forewing broader; cell cu, higher; surface spinules present or absent. Male paramere 
broad; female términalia différent 2 cee en too 109 
Large species, forewing longer than 2.65 mm. Surface spinules of forewing always present. Terminalia as in Figs 
118-119 OM OUCH CUS ERE O ROR OL OT RE TOTO Trioza remota 
Small species, forewing shorter than 2.6 mm. Surface spinules present or absent. Terminalia different. On Amaran- 
thaceaen RE Sc Ose oa e Trioza chenopodii 
Body green or yellow: mne Re 111 
Body colour différent... ii ER TR I RC eT 113 
Antennal segments 4-6 yellow or light ochreous; apical two or three dark segments strongly contrasting with the 
lshtenmmorebasalisegmentsAON ADIAC CA Cm" Trioza apicalis 
Antennal segments 4-6 dark ochreous or light brown; antennal segments gradually becoming darker from segment 
3 ABER. ern ANREISE Re Re RER 112 


Paramere with short dorsal projection and narrow anterior lobe (Fig. 120). Female subgenital plate with terminal 
process forming a long parallel sided projection (Fig. 121). On Anthriscus and other Apiaceae ... Trioza anthrisci 


113 


114 


115 


116 


117 


118 


119 


126 


D 
(09) 


Jumping plant-lice of Belarus 


Paramere with long dorsal projection and narrow anterior lobe (Fig. 122). Female subgenital plate with short and 


repularlygtaperngatermimaliprocessi(Kio #28) O NUASEH PIUME ee Trioza laserpitii 
SUL AC GomMulles presti in Ghee! cal AT TORENT ee 114 
Sunfacesspinulesgabsentgorgpresen GL NOS NICE CURE TERE ORE IRE etero 120 
Surface}spinulesfabsent from celllr OP ONE WIN pe" MN Nes Bactericera maura 
SUBLACEKSPINUESIPIESCAMINICENTAO OTE WIN DE seen: 115 
Aneamall sgamens d-7/ ALE NI ne ne nere aprira 116 
Atom scans 457 Mn once ee ere Re NT EEE TR PU 117 


Surface spinules entirely covering cell r of forewing. Abdominal venter whitish, distinctly lighter than dorsum. 
Male paramere, in profile, sinuous with wide base and slender apex (Fig. 124). Ventral margin of female subgenital 
Dates ap hUOHCONCAVEISUDAP1CAllyI ONISA Eee Bactericera albiventris 
Surface spinules of forewing present only in basal half of cell r,. Abdominal venter not whitish, or the same colour 
as dorsum. Male paramere, in profile, straight, with mostly subparal margins (Fig. 125). Ventral margin of female 


subgenital plate straight or convex subapically. On Salix. ......... i Bactericera salicivora 
Surface spinules in apical half of forewing covering the whole surface up to the veins; wing angular apically (Fig. 
2) MÉminala SNS 227 ONCOMAMMIDANSTEE ne: Bactericera acutipennis 
Surface spinules leaving spinule-free stripes along veins of forewing ................. 118 
Antennal segments 4 and 6 with each a group of rhinaria, therefore strongly dilated apically. Terminalia as in Figs 
2 IZORONA CHEN APRE Rice cose oa kc at ri Bactericera femoralis 
Antennal segments 4 and 6 with each | rhinarium, not strongly dilated apically.................. ii 119 
Surface spinules of forewing also present in distal half of cell r, and entirely covering cell c+sc. On Potentilla 
DOSE SAAS IATA IRIS ARA AI TOI ITTOTE Bactericera reuteri 
Surface spinules of forewing present only in proximal half of cell r, and strongly reduced in cell c+sc. On Geum 
SANTANA IRA III DI SOIT dae ENG Sabdoabubeusscevavenesneces Bactericera bohemica 
Genal processes at most as long as two thirds of vertex. Antenna entirely dark brown to black. Terminalia as in Figs 
1802 Oly phagousronidicotyledonousiherbSpessesesereotere ecco eee eens Bactericera nigricornis 
Genal processes about as long as vertex. At least antennal segment 3 light... 121 
ANIME, COMO DNV Were, Cid ZV ATAENT Cloc0<00000000000000000000000600070000006005006000063000000000000000000000000000 Bactericera calcarata 
At |kast anal SEINE E sect RIT IAT RR LE sic 122 
Antennal segments 3-5 partly or entirely light. On Sanguisorba ............. iii Bactericera modesta 
ANTEIMA ESCO men tesa ote Aran eon AKI N ENT e entre 123 
Borewineswides im tine manatee, Oln BAALEAIS WKY 2171S coc00c00cc0200000000000000000000000000000000000000000000600000000000000000 Trioza scottii 
KorewineswidesingapticalithirdSKONESa 3 o0cec05000000006600000060050000080000090000000055000005000000 e nnt 124 
HOrEWIN PAWI LAVE INPNENTITE MAT ATK ee Bactericera curvatinervis 
DarCo OUHONVEIN AT tel Ore WIN LITE AU CEI MOXAS DO 125 
Dark spot on vein A of forewing distinct. Male paramere, in profile, more or less straight with large anteriorly 
directed tooth (Fig. 132). Female proctiger short with apical two thirds of dorsal outline strongly convex ............ 
ns ann TEATINO n tant de suctBisbcidsceiFasascsuusendebioaeavaseases Bactericera striola 
Dark spot on vein A of forewing indistinct. Male paramere sickle-shaped or with small apical hook. Female proc- 
tiger longer with apical two thirds of dorsal outline straight or CONCAVE .......................... ii 126 
Male parameres, in profile, sickle-shaped. Female subgenital plate long with indistinct, long apical process ........ 
5000006600800500000030000 NEED API ARE ne SESE EE RE ER ET RTE PE RE ICE AIAR OUT EP EEE EE FETE EE Bactericera substriola 


Male parameres, in profile, straight, lamellar, with small apical tooth. Female subgenital plate short with distinct 
Shorteapicalg process TREE a Bactericera parastriola 


24 L. Serbina, D. Burckhardt & O. Borodin 


Figs 1-10. (1) Cacopsylla pyricola, forewing. (2) Bactericera acutipennis, forewing. (3) Psyllopsis fraxinicola, male terminalia. (4) 
Psyllopsis fraxinicola, female terminalia. (5) Psyllopsis discrepans, forewing. (6) Psyllopsis discrepans, paramere. (7) 
Psyllopsis discrepans, female terminalia. (8) Psyllopsis distinguenda, forewing. (9) Psyllopsis distinguenda, paramere. (10) 
Psyllopsis distinguenda, female terminalia. 


Jumping plant-lice of Belarus 25 


Figs 11-18. (11) Psyllopsis fraxini, forewing. (12) Psyllopsis fraxini, paramere. (13) Psyllopsis fraxini, female terminalia. (14) 
Strophingia ericae, male terminalia. (15) Aphalara exilis, forewing surface spinules. (16) Aphalara ulicis, forewing 
surface spinules. (17) Aphalara exilis, paramere. (18) Aphalara ulicis, paramere. 


26 L. Serbina, D. Burckhardt & O. Borodin 


Figs 19-29. (19) Aphalara longicaudata, forewing surface spinules. (20) Aphalara longicaudata, paramere. (21) Aphalara 
longicaudata, female terminalia. (22) Aphalara calthae, forewing surface spinules. (23) Aphalara polygoni, forewing 
surface spinules. (24) Aphalara borealis, forewing surface spinules. (25) Aphalara avicularis, forewing surface spinules. 
(26) Aphalara freji, forewing surface spinules. (27) Aphalara borealis, male terminalia. (28) Aphalara borealis, aedeagus. 
(29) Aphalara borealis, female terminalia. 


Jumping plant-lice of Belarus 


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Figs 30-40. (30) Aphalara avicularis, male terminalia. (31) Aphalara avicularis, aedeagus. (32) Aphalara avicularis, female 
terminalia. (33) Aphalara freji, male terminalia. (34) Aphalara freji, aedeagus. (35) Aphalara freji, female terminalia. 
(36) Craspedolepta artemisiae, forewing surface spinules. (37) Craspedolepta latior, forewing surface spinules. (38) 


Craspedolepta malachitica, forewing surface spinules. (39) Craspedolepta latior, forewing. (40) Craspedolepta 
malachitica, forewing. 


28 L. Serbina, D. Burckhardt & O. Borodin 


Crt 


Figs 41-48. (41) Craspedolepta artemisiae, male terminalia. (42) Craspedolepta artemisiae, female terminalia. (43) Craspedolepta 
latior, male terminalia. (44) Craspedolepta latior, female terminalia. (45) Craspedolepta malachitica, male terminalia. 
(46) Craspedolepta malachitica, female terminalia. (47) Craspedolepta subpunctata, paramere. (48) Craspedolepta 
subpunctata, female terminalia. 


Jumping plant-lice of Belarus DI 


Figs 49-58. (49) Craspedolepta innoxia, forewing surface spinules. (50) Craspedolepta innoxia, paramere. (51) Craspedolepta 
innoxia, female terminalia. (52) Craspedolepta omissa, forewing surface spinules. (53) Craspedolepta omissa, paramere. 
(54) Craspedolepta omissa, female terminalia. (55) Craspedolepta nebulosa, forewing. (56) Craspedolepta nebulosa, 
male terminalia. (57) Craspedolepta nebulosa, paramere. (58) Craspedolepta nebulosa, female terminalia. 


30 L. Serbina, D. Burckhardt & O. Borodin 


Figs 59-66. (59) Craspedolepta nervosa, forewing. (60) Craspedolepta bulgarica, forewing. (61) Craspedolepta nervosa, male 
terminalia. (62) Craspedolepta nervosa, female terminalia. (63) Craspedolepta bulgarica, male terminalia. (64) 
Craspedolepta bulgarica, female terminalia. (65) Craspedolepta crispati, male terminalia. (66) Craspedolepta sonchi, 
male terminalia. 


Jumping plant-lice of Belarus Bil 


N 


Figs 67-74. (67) Craspedolepta crispati, female terminalia. (68) Craspedolepta sonchi, female terminalia. (69) Livilla horvathi, 
forewing. (70) Livilla radiata, forewing. (71) Cacopsylla breviantennata, forewing. (72) Cacopsylla breviantennata, male 
terminalia. (73) Cacopsylla pruni, forewing. (74) Cacopsylla pruni, male terminalia. 


32 L. Serbina, D. Burckhardt & O. Borodin 


og RIM 


Figs 75-82. (75) Cacopsylla breviantennata, female terminalia. (76) Cacopsylla pruni, female terminalia. (77) Cacopsylla pyrisuga, 
male terminalia. (78) Cacopsylla pyrisuga, female terminalia. (79) Cacopsylla picta, male terminalia. (80) Cacopsylla 
picta, female terminalia. (81) Cacopsylla melanoneura, male terminalia. (82) Cacopsylla affinis, male terminalia. 


Jumping plant-lice of Belarus 33 


Figs 83-90. (83) Cacopsylla albipes, forewing. (84) Cacopsylla crataegi, forewing. (85) Cacopsylla pyri, male terminalia. (86) 
Cacopsylla pyri, female terminalia. (87) Cacopsylla pyricola, male terminalia. (88) Cacopsylla pyricola, female terminalia. 
(89) Cacopsylla zetterstedti, male terminalia. (90) Cacopsylla hippophaes, male terminalia. 


34 L. Serbina, D. Burckhardt & O. Borodin 


Figs 91-97. (91) Cacopsylla zetterstedti, female terminalia. (92) Cacopsylla hippophaes, female terminalia. (93) Cacopsylla 
brunneipennis, male terminalia. (94) Cacopsylla nigrita, male terminalia. (95) Trichochermes walkeri, forewing. (96) 
Trioza centranthi, male terminalia. (97) Trioza centranthi, female terminalia. 


Jumping plant-lice of Belarus 35 


Figs 98-105. (98) Trioza galii, male terminalia. (99) Trioza galii, female terminalia. (100) Trioza velutina, male terminalia. (101) 
Trioza velutina, female terminalia. (102) Trioza urticae, male terminalia. (103) Trioza urticae, female terminalia. (104) 
Trioza proxima, male terminalia. (105) Trioza proxima, female terminalia. 


36 L. Serbina, D. Burckhardt & O. Borodin 


Figs 106-112. (106) Trioza dispar, male terminalia. (107) Trioza dispar, female terminalia. (108) Trioza tatrensis, male terminalia. 
(109) Trioza tatrensis, female terminalia. (110) Trioza rotundata, male terminalia. (111) Trioza rotundata, female 
terminalia. (112) Trioza cerastii, male terminalia. 


Jumping plant-lice of Belarus 3)7/ 


Figs 113-119. (113) Trioza cirsii, male terminalia. (114) Trioza cirsii, female terminalia. (115) Trioza abdominalis, male terminalia. 
(116) Trioza alacris, male terminalia. (117) Trioza alacris, female terminalia. (118) Trioza remota, male terminalia. 
(119) Trioza remota, female terminalia. 


38 L. Serbina, D. Burckhardt & O. Borodin 


Figs 120-125. (120) Trioza anthrisci, male terminalia. (121) Trioza anthrisci, female terminalia. (122) Trioza laserpitii, male terminalia. 
(123) Trioza laserpitii, female terminalia. (124) Bactericera albiventris, male terminalia. (125) Bactericera salicivora, 
male terminalia. 


Jumping plant-lice of Belarus 39 


Figs 126-132. (126) Bactericera acutipennis, male terminalia. (127) Bactericera acutipennis, female terminalia. (128) Bactericera 
femoralis, male terminalia. (129) Bactericera femoralis, female terminalia. (130) Bactericera nigricornis, male 
terminalia. (131) Bactericera nigricornis, female terminalia. (132) Bactericera striola, male terminalia. 


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L. Serbina, D. Burckhardt & O. Borodin 


44 


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Revue suisse de Zoologie (March 2015) 122 (1): 45-54 


ISSN 0035-418 


Three new rotundabaloghid mites (Acari: Uropodina) from Hong Kong 


Jeno Kontschan 


Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of Sciences, H-1525 Budapest, PO. Box 


102, Hungary. E-mail: kontschan.jeno@agrar.mta.hu 


Abstract: Three new species of the family Rotundabaloghidae are described from Hong Kong. Angulobaloghia staryi sp. 
nov. differs from the other Angulobaloghia Hirschmann, 1979 species in the shape and ornamentation of the genital shield 
of the female. Rotundabaloghia (Rotundabaloghia) hongkongensis sp. nov. has three pairs of short setae (Stl, V2 and 
V6) on the ventral idiosoma, which is unique in the subgenus Rotundabaloghia (Rotundabaloghia) Hirschmann, 1975a. 
The long, robust and curved setae in the big ventral cavity of Depressorotunda (Depressorotunda) taurina sp. nov. 1s a 
character so far unknown in the subgenus Depressorotunda (Depressorotunda) Kontschan, 2010a. 


Keywords: East Asia - taxonomy - turtle-mites. 


INTRODUCTION 


The rotundabalogid mites are small, rounded, soil 
dwelling members of the suborder Uropodina, which have 
a typical circumtropical distribution. Rotundabaloghids 
can be found in South and Central America, Africa, 
South-East Asia and the Austral-Asian region 
(Kontschan, 2010b). Three taxa of the family [the genus 
Angulobaloghia Hirschmann, 1979 and the subgenera 
Rotundabaloghia (Rotundabaloghia) Hirschmann, 1975a 
and Depressorotunda (Depressorotunda) Kontschän, 
2010a] occur only in the South-East Asian and Austral- 
Asian regions (Kontschan, 2010b, 2011; Kontschan & 
Stary, 2012). Several parts of these regions have been 
intensively investigated, but there are some areas from 
where no rotundabaloghid mites are known. 

I spent some days in the Arachnida collection of the 
Natural History Museum in Geneva, where I separated 
several rotundabalogid mites. Here I present my first 
results of this visit, with the descriptions of three new 
species from Hong Kong. 


MATERIAL AND METHODS 


Specimens were cleared in lactic acid and drawings were 
made with the aid of a drawing tube. All specimens are 
stored in ethanol and deposited in the Natural History 
Museum in Geneva (MHNG). The nomenclature of the 
ventral setation, and the generic and subgeneric systems 
with the diagnoses of these taxa follow Kontschan 
(2010b). The chelicerae of Rotundabaloghidae are often 
hidden inside the body and in that case they cannot be 
observed without breaking the specimens. However, the 


Manuscript accepted 24.09.2014 
DOI: 10.5281/zenodo.14581 


chelicerae do not have important characters for species 
identification within this family (chelicerae have a 
long and apically rounded fixed digit, and the internal 
sclerotized node is visible). Therefore, this character was 
not always studied in order not to cause undue damage 
to the type specimens. For the examined material, the 
original label data in French are given, this to avoid 
incorrect interpretations. 


Abbreviations: h = hypostomal setae, St = sternal 
setae, im = internal malae, V = ventral setae. All 
measurements and the scales in the figures are given in 
micrometres (um). 


TAXONOMY 


Angulobaloghia staryi sp. nov. 
Figs 1-11 


Holotype: MHNG, sample SBH-96/19; female; Hong 
Kong (New Territories), Tai Mo Shan Country Park, 
barrier ravine, petits îlots de forêt, prélèvement de sol 
entre les racines de Machilus sp. (Lauraceae), 750 m; 
9.XII.1996; leg. B. Hauser (extraction par appareil 
Berlese a Genève). 


Paratypes: MHNG; | female, 5 males; same data as for 
holotype. 


Diagnosis: Genital shield subtriangular and its surface 
with oval pits. Setae V7 and V8 smooth and needle- 
like, situated near end of pedofossae IV. Setae on 
dorsal side of body smooth. The new species differs 
from the other known Angulobaloghia species by the 
shape (subtriangular and with wide basis) and pit-like 


46 J. Kontschan 


ornamentation of the genital shield in females and by 
the central position of setae V6 in both sexes. 


Description of female: Length of idiosoma 370- 
380 um, width 340-350 um (n=2). Shape circular, 
posterior margin rounded, color reddish brown. 

Dorsal idiosoma (Fig. 1): Marginal and dorsal shields 
fused. Dorsal setae basally curved, margins smooth 
(about 17-23 um long) (Figs 2-3). Surface with small 
oval pits (Figs 2-3). 

Ventral idiosoma (Fig. 4): Sternal and ventral shields 
without sculptural pattern, but a linguliform, shallow 
depression present near setae Stl. All sternal setae 
smooth, needle-like and short (about 5-6 um long), Stl 
situated at level of central region of coxae II, St2 at level 


of posterior margin of coxae II, St3 at level of posterior 
margin of coxae III, St4 at level of anterior margins of 
coxae IV, St5 situated at level of pedofossae IV. Ventral 
setae smooth and needle-like. V2 about 6-7 um long, 
situated near basal line of genital shield. V7 and V8 
about 15-17 um long, situated near end of pedofossae IV. 
Setae ad similar in shape and length to V2 setae, situated 
laterally to anal opening. V6 setae absent. One pair of 
lyrifissures situated anterior to Stl, one pair near St4, 
and two pairs of other sensory organs plus one pair of 
lyrifissures near V7 and V8. Stigmata situated between 
coxae II and III. Peritremes hook-shaped. Genital shield 
long and subtriangular, its surface with oval pits and its 
apical margin rounded. Base of tritosternum narrow, vase- 
like, tritosternal laciniae smooth, subdivided into three 


Figs 1-5. Angulobaloghia staryi sp. nov., female holotype. (1) Body in dorsal view. (2) Setae and ornamentation in central area of 
dorsal shield. (3) Setae and ornamentation in lateral part of dorsal shield. (4) Body in ventral view. (5) Ventral setae. 


New rotundabaloghid mites from Hong Kong 47 


smooth branches in its distal half (Fig. 6). Pedofossae 
deep, their surface smooth, separated furrows for tarsi IV 
present. 

Gnathosoma (Fig. 6): Corniculi horn-like, internal malae 
smooth and longer than corniculi. Hypostomal setae as 


follows: hl long (about 22-23 um), smooth and needle- 
like, h2 and h3 short (about 6-7 um), smooth and needle- 
like, h4 marginally serrate and short (about 5-6 um). 
Apical part of epistome marginally pilose. Palp with 
smooth setae. Chelicerae not clearly visible. 


Figs 6-11. Angulobaloghia staryi sp. nov., female holotype (6-10) and male paratype (11). (6) Ventral view of gnathosoma, tritosternum 
and coxae I. (7) Ventral view of leg I. (8) Ventral view of leg II. (9) Ventral view of leg III. (10) Ventral view of leg IV. (11) 
Ventral view of intercoxal area of male. 


48 


J. Kontschan 


Key to the species of Angulobaloghia (modified, after Kontschän, 2010b, 2011): 


| ventral;shield’ornamentedee.......22. 2er TE A. aokii (Hiramatsu, 1977) 
- ventral;shield:smooth.... 2.222. een rennen ee SERIORNEEEENEEEDEEREE 2 
? genitaltshicl dlofetemaleysemi=cinc ul ARE A. danyii (Kontschan, 2008a) 
= genital:shield’offemaleitriangular.....e..22.2022022° 12er TE EREEENIR 3 
3 peritremes mushroom-shaped...-... 2.2832 ca n, 4 
— peritremesihook=shaped.....=.......ue RI RR ne 7 
4 genitalishield:withpatten star nee consent ee OE 5 
_ penital‘shieldiwithout:pattermn 52e AR NE ee cea ee cee ee 6 
5 StSithree timesilon2enthaNn tONENStern als ELA ee ER 

CECI IAA ER RER LE A. cuyi (Hiramatsu & Hirschmann in Hirschmann & Hiramatsu, 1992) 
_ St3 as long as other sternal setae........................... A. pyrigynella (Hirschmann in Hirschmann & Hiramatsu, 1992) 
6 sternalsshieldayuthtornamentallon ee A. vietnamensis (Kontschän, 2008a) 
= sternal shield without ornamentation .…............................................................ A. triangulata (Kontschan, 2008a) 
7 sternal'shield:with’ornamentationi............rcrr rit nn 8 
= sternal’shield’without ornamentation’ 22 en eee n RERRERESIER 9 
8 SD Graal SiS} Worse LINE SION p erdtha ISU E NT A. indica Kontschan, 2011 
_ St2fand/St3fasilongiasStls-st-® A. luzonensis (Hiramatsu & Hirschmann in Hirschmann & Hiramatsu, 1992) 
9 genital Shield'Withiormamentation Sim ana it RN n 10 

senital shield without ornamentation....... sense res ee eae ne 12 
10 anterior margin of genital shield smooth and rounded, setae V6 smooth ......................\ i 11 
- anterior margin of genital shield serrate, setae V6 with one pair of lateral spines........................... 2200 

ini ET e VOICE RATORI ON TRO MOR TT A. scrobia Kontschän & Stary, 2011 
11 basal part of genital shield narrow, V6 situated near V7 and V8 ................ A. angustigynella (Hirschmann, 1975b) 


= basal part of genital shield wide, V6 situated in central area of ventral shield ............................... A. staryi Sp. nov. 


1 St2 and St3 three times longer than Stl....................... 


= St2 and St3 as long as Stl 


_ additional setae on sternal shield absent 


Legs (Figs 7-10). All legs with ambulacral claws and 
smooth and needle-like setae. 


Description of male: Length of idiosoma 370-410 um, 
width 340-370 um (n=5). 

Dorsal idiosoma: Ornamentation and chaetotaxy of 
dorsal shield as for female. 

Ventral idiosoma (Fig. 11): Four pairs of sternal setae 
situated anterior and one pair posterior to genital shield, 
these setae smooth, needle-like and about 8-10 um long 
(except St3 being about 14 um). St1 situated near anterior 
margin of sternal shield, St2 at level of posterior margin 
of coxae II, St3 at level of posterior margin of coxae 
III, St4 near anterior margin of genital shield, St5 near 
posterior margin of genital shield. One pair of lyrifissures 
situated near Stl and one pair near St4. Surface of sternal 
shield with numerous oval pits anterior to genital shield. 
Surface of ventral shield, and shape and size of ventral 
setae as in female. Genital shield oval and situated 
between coxae IV. 

Larva and nymphs unknown. 


Etymology: I dedicate the new species to my colleague 
and dear friend, to Dr. Josef Stary (Biology Centre AS 
CR, Institute of Soil Biology, Ceské Budèjovice, Czech 
Republic). 


13 additional setae on sternal shield present .................... 


SIOE ennemies A. angullogynella (Hirschmann, 1975b) 
LR ESI ER ETORN 13 
RNA di ico ou A. latigynella (Hirschmann, 1975b) 
PRI e I e A. tamilica Kontschan, 2011 


Rotundabaloghia (Rotundabaloghia) hongkongensis 
Sp. nov. 
Figs 12-20 


Holotype: MHNG, sample SBH-96/19; female; Hong 
Kong (New Territories), Tai Mo Shan Country Park, 
barrier ravine, petits îlots de forét, prélèvement de sol 
entre les racines de Machilus sp. (Lauraceae), 750 m; 
9.XII.1996; leg. B. Hauser (extraction par appareil 
Berlese à Genève). 


Paratypes: MHNG; 6 females, 5 males; same data as 
for holotype. 


Diagnosis: Genital shield linguliform and its surface 
with irregular pits. Setae V6 and V8 shorter than V2 and 
V7. Dorsal setae on caudal area wide and marginally 
pilose. Setae Stl very short, similar in shape and 
length to St4. Rotundabaloghia (Rotundabaloghia) 
hongkongensis sp. nov. has three pairs of short setae 
(Stl, V2 and V6) on the ventral part of its body, 
which is a unique character within the subgenus 
Rotundabaloghia (Rotundabaloghia) Hirschmann, 
1975a. 


Description of female: Length of idiosoma 370-380 
um, width 340-350 um (n=2). Shape of idiosoma 
circular, posterior margin rounded, color reddish brown. 


New rotundabaloghid mites from Hong Kong 49 


Dorsal idiosoma (Fig. 12): Marginal and dorsal shields 
fused. Three pairs of setae in row j short, smooth and 
needle-like (about 12-13 um), other setae on dorsal 
idiosoma slightly pilose (Fig. 13) (about 27-30 um), setae 
in caudal area of dorsal part wide and spatuliform (about 
30-31 um). Dorsal idiosoma with oval pits (Fig. 13). 

Ventral idiosoma (Fig. 14): Sternal and ventral shields 
with some oval pits. Setae St1 smooth, short (about 8 um) 
and needle-like, situated near anterior margin of sternal 


shield, St2 smooth and long (about 29 um), situated at 
level of posterior margin of coxae II, St3 smooth and 
long (about 31 um), at level of central area of coxae III, 
St4 similar in shape and length to Stl, at level of anterior 
margins of coxae IV. St5 absent. Ventral setae smooth and 
needle-like. V2 and V6 situated near posterior margin of 
coxae IV, V2 long (about 44-46 um), V6 short (about 13- 
14 um). V8 short (about 14-15 um) and situated near end 
of pedofossae IV. V7 long (about 35-37 um) and situated 


Figs 12-16. Rotundabaloghia (Rotundabaloghia) hongkongensis sp. nov., female holotype (12-15) and male paratype (16). (12) 
Body in dorsal view. (13) Dorsal setae and ornamentation. (14) Ventral view of body. (15) Ventral view of gnathosoma, 
tritosternum and coxae I. (16) Ventral view of intercoxal area of male. 


50 J. Kontschan 


between V8 and ad. Setae ad similar in shape and length 
to V6, lateral to anal opening. Stigmata situated between 
coxae II and III. Peritremes hook-shaped. Genital shield 
long and linguliform, surface with irregular pits, its apical 
margin rounded. Base of tritosternum narrow, tritosternal 
laciniae smooth, subdivided into three smooth branches 
(Fig. 15). Pedofossae deep, their surface smooth, 
separated furrows for tarsi IV present. 

Gnathosoma (Fig. 15): Corniculi horn-like, internal 
malae longer than corniculi and smooth. Labrum apically 
pilose. Hypostomal setae as follows: hl long (about 
27 um), smooth and needle-like, h2 (about 13 um), h3 and 
h4 short (about 6-7 um), smooth and needle-like. Apical 
part of epistome slightly pilose. Palp with smooth setae 
except for one slightly serrate seta on palp trochanter. 
Chelicerae not clearly visible. 

Legs (Figs 17-20). All legs with ambulacral claws and 
smooth and needle-like setae, tarsi III and IV with more 
robust setae than other articles. 


Description of male: Length of idiosoma 370-410 um, 
width 340-370 um (n=5). 

Dorsal idiosoma: Ornamentation and chaetotaxy of 
dorsal shield as for female. 


Ventral idiosoma (Fig. 16): Four pairs of sternal setae 
situated anterior to genital shield, these setae smooth, 
needle-like. Stl, St4 and St5 short (about 7-9 um), St2 
and St3 long (about 29-30 um). Stl situated near anterior 
margin of sternal shield, St2 situated at level of posterior 
margin of coxae II, St3 at level of central area of coxae 
III, near anterior margin of genital shield, St5 near lateral 
margin of genital shield. Surface of sternal shield with 
numerous oval pits. Surface of ventral shield, and shape 
and size of ventral setae as in female. Genital shield 
circular and situated between coxae IV. 

Larva and nymphs unknown. 


Etymology: The name of the new species refers to the 
city (Hong Kong) where it was collected. 


Depressorotunda (Depressorotunda) taurina sp. nov. 
Figs 21-29 


Holotype: MHNG, sample SBH-96/19; female; Hong 
Kong (New Territories), Tai Mo Shan Country Park, 
barrier ravine, petits îlots de forêt, prélèvement de sol 
entre les racines de Machilus sp. (Lauraceae), 750 m; 


1 


Figs 17-20. Rotundabaloghia (Rotundabaloghia) hongkongensis sp. nov., female holotype. (17) Ventral view of leg I. (18) Ventral 
view of leg II. (19) Ventral view of leg III. (20) Ventral view of leg IV. 


New rotundabaloghid mites from Hong Kong SI 


Key to the species of Rotundabaloghia (Rotundabaloghia) species (modified, after Kontschan, 2010b): 


I IVISISCIACISTA © Oth RR EEE EEE EEE EEEEEENER 2 
= \VISISETA CIN OSIMO LP e e SOI INI EEE RE EE PRET tai 7 
À WVSISnOrterginantotneraVentra SE AE n renna e nine een sen One ie er nnn ne den rater neto cernes ones sen a detente 5 
— VS. lon as V2 OL AINA AE e I Ot ottieni cinici tralicci 3 
3 Val ONE ersthantotherav en na IIS E A R. (R.) kaszabi Hirschmann, 1975b 
- VARSTIONLEAS ONE ENTIER nn ennemie nd Ness eee nr etes sne tree encens asc eee 4 
4 Onmnamentavionybetwiecnevs/sPleSel ER ee R. (R.) makilingoides Hirschmann & Hiramatsu, 1992 
- OHMAMENTAONIDELWE CN TEA DS EN re R. (R.). makilingensis Hirschmann & Hiramatsu, 1992 
5 IVCTASI ON DAS rene drone A eee rh R. (R.) hongkongensis sp. nov. 
- IV OB ON SCIA NANNI PRE RR AR Ii init EE TE OPEN ina 6 
6 WSEAS] OM De EI ECA ARRE IE MEIN LT INR RT R. (R.) korsosi Kontschän, 2008b 
= VS otre CA RP vouensdeasieess R. (R.) hirschmanni Hiramatsu, 1977 
7 SETACIACIPIOSCR ZERI O ce chose Gace COASTS CEE CeO CPR Cee eRe R. (R.) baloghi Hirschmann, 1975b 
- SEES AA TITAN sonen PO es sect 8 
8 W/7/ 0 | OS CHE N nti deci 9 
— \VAZASIN OO (RR RETE oe ER tenant ue se ettse té tetes es 10 
9 SulisSt2fandiStsimuchmilongerd((1(0$) ith ants APE R. (R.) macroseta Hirschmann, 1975b 


- Stl, St2 and St3 not much longer (4x) than St4.......... 
10 Stl longer and wider than St2 and St3 ........................ 


- Stl not longer and wider than St2 and St3 


11 VPN OIIMESION LE MNANATE PP 
— VERBOTEN er 


9.XII.1996; leg. B. Hauser (extraction par appareil 
Berlese à Genève). 


Paratype: MHNG; 1 female; same data as for holotype. 


Diagnosis: Ventral cavity quadrangular, with one pair 
of long, robust and curved setae. Genital shield lingu- 
liform, its surface with shallow pits. Three pairs of short 
ventral setae situated near lateral margins of ventral ca- 
vity. The long, robust and curved setae in the big ventral 
cavity in Depressorotunda (Depressorotunda) taurina 
Sp. nov. is a previously not observed character with- 
in the subgenus Depressorotunda (Depressorotunda) 
Kontschan, 2010a. 


Description of female: Length of idiosoma 290-310 
um, width 260-270 um (n=2). Shape circular, posterior 
margin rounded, color reddish brown. 

Dorsal idiosoma (Fig. 21): Marginal and dorsal shields 
fused. Dorsal setae long (about 27-33 um long) and 
smooth (Figs 22-23), dorsal idiosoma with small oval 
pits (Figs 22-23). 

Ventral idiosoma (Fig. 24): Sternal and ventral shields 
without sculptural pattern. All sternal setae smooth, 
needle-like and short (about 7-9 um), St! situated at level 
of anterior margin of coxae II, St2 at level of posterior 
margin of coxae II, St3 at level of posterior margin of 
coxae III, St4 near basal edges of genital shield. Dorsal 
cavity large, quadrangular, wider (about 75-76 um) than 
long (about 34-35 um). Ventral setae smooth and needle- 
like. V1 placed in ventral cavity, long (about 34-35 um), 
robust and curved. V2, V3 and V4 short (about 9-10 um) 


Sa Te SERRE R. (R.) mahunkai Hirschmann, 1975b 
NEO o R. (R.) monomacroseta Hirschmann, 1975b 
III II TT 11 
PIE RI RA 0 R. (R.) kaszabisimilis Hirschmann, 1975b 
RR RI En ete R. (R.) pilosa Hirschmann, 1975b 


and needle-like. V2 situated near anterior edges of ventral 
cavity, V3 and V4 near basal edges of ventral cavity 
(Fig. 25). Adanal setae absent. One pair of lyrifissures 
situated near St4. Stigmata situated between coxae II 
and III. Peritremes hook-shaped. Genital shield long and 
linguliform, surface with shallow pits, its apical margin 
rounded. Tritosternum covered by coxae I (Fig. 26), its 
parts not visible. Pedofossae deep, their surface smooth, 
separated furrows for tarsi IV present. 

Gnathosoma (Fig. 26): Corniculi horn-like, internal 
malae shorter than corniculi and smooth. Hypostomal 
setae as follows: hl long (about 27 um), smooth and 
needle-like, h2, h3 and h4 short (about 12-15 um), 
smooth and needle-like. Apical part of epistome weakly 
pilose. Chelicerae not clearly visible. 

Legs (Figs 26-29). All legs with ambulacral claws and 
smooth, needle-like setae. 

Male, larva and nymphs unknown. 


Etymology: The new species is named after the long, 
robust and curved V1 setae, which are similar to bull 
horns; “taurina” is the Latinized adjective of “taurus” (= 
bull). 


ACKNOWLEDGEMENTS 


I am very grateful to Dr. Peter Schwendinger (MHNG) 
for his kind hospitality during my stay in Geneva, and 
to Dr. Jerzy Bloszyk (University of Poznan, Poland) for 
commenting on an earlier version of the manuscript. 


52 J. Kontschan 


Figs 21-25. Depressorotunda (Depressorotunda) taurina sp. nov., female holotype. (21) Body in dorsal view. (22) Setae and 
ornamentation in central area of dorsal shield. (23) Setae and ornamentation in lateral part of dorsal shield. (24) Body in 
ventral view. (25) Intercoxal area and ventral cavity in ventral view. 


New rotundabaloghid mites from Hong Kong 53 


Me 


-d 


Figs 26-29. Depressorotunda (Depressorotunda) taurina sp. nov., female holotype. (26) Ventral view of leg I and gnathosoma. (27) 
Ventral view of leg II. (28) Ventral view of leg III. (29) Ventral view of leg IV. 


54 J. Kontschän 


Key to the species of Depressorotunda (Depressorotunda) (modified, after Kontschan, 2010b, Kontschan & Stary, 


2012): 

l genital shield of female and sternal shield of male ornamented with oval pits.…............................................ 
VILLAR III SIR E ARE REN IRE NOA ACER RS EA E ene D. (D.) alveolata Kontschan & Stary, 2011 

- genital shield of female and sternal shield of male without ornamentation ...................... 2 

2 all veritral setae ‘smooth... Rss he O I O TT 3 


- ONEIPAIMONVEN TAlISC LAC) DOS CPE 


RARA I D. (D.) batuensis Kontschan & Stary, 2012 
peritreme R-shaped, posteriormost dorsal setae conspicuously longer than those in anterior position ..................... 
STATO ee D. (D.) seticaudata Kontschan, 2010a 
peutremehook-shapedsmostidorsalisetacie qua limi) en SN EE 4 


RE ete D. (D.) mirifica Kontschan, 2010a 
a Pe Oe Te ne D. (D.) taurina sp. nov. 


ventral cavity of male conspicuously enlarged, reaching sternal region, with genital shield in medial part; most 
he er See ee ee D. (D.) malayana Kontschan, 2010a 


— ventral cavity of male smaller, not reaching sternal region, genital shield situated outside cavity; most dorsal setae 


3 
4 WS} BIS NOTE BS WD ANNEE 
— V3 two times longer than V2 and V4........................ 
5 ventral cavity wider than long (w:1 =1:0.4 ................. 
— ventral cavity longer than wide (w:1 = 1:2.................. 
6 
dorsale ta WITNIAIStA lIDITO SILVER 
SMOOME Rn ere ree ee 
REFERENCES 


Hiramatsu N. 1977. Gangsystematik der Parasitiformes. Teil 
239. Teilgang einer neuen Rotundabaloghia-Art aus Japan 
(Dinychini, Uropodinae). Acarologie, Schriftenreihe für 
Vergleichende Milbenkunde 23: 19-20. 

Hirschmann W. 1975a. Gangsystematik der Parasitiformes. Teil 
201. Die Gattung Rotundabaloghia nov. gen. Hirschmann 
1975 (Dinychini, Uropodinae). Acarologie, Schriftenreihe 
fiir Vergleichende Milbenkunde 21: 23-26. 

Hirschmann W. 1975b. Gangsystematik der Parasitiformes. Teil 
203. Teilgänge, Stadien von 16 neuen Rotundabaloghia- 
Arten (Dinychini, Uropodinae). Acarologie, Schriftenreihe 
für Vergleichende Milbenkunde 21: 28-34. 

Hirschmann W. 1979. Stadiensystematik der Parasitiformes I. 
Stadienfamilien und Stadiengattungen der Atrichopygidiina, 
erstellt im Vergleich zum Gangsystem Hirschmann 1979. 
Acarologie, Schriftenreihe für Vergleichende Milbenkunde 
26: 75-70. 

Hirschmann W., Hiramatsu N. 1992. 34 Rotundabaloghia- 
Arten aus Asien (Japan, Neuguinea, Philippinen, Borneo) 
(Dinychini, Uropodinae). Acarologie, Schriftenreihe fiir 
Vergleichende Milbenkunde 39: 9-25. 


Pe eee RTE SM NE D. (D.) thailandica Kontschan, 2010a 


Kontschan J. 2008a. New and rare Rotundabaloghia species 
(Acari: Uropodina) from the tropics. Opuscula Zoologica 
(Budapest) 38: 15-41. 

Kontschan J. 2008b. Rotundabaloghia korsosi sp. nov. (Acari: 
Uropodina) from Taiwan. Collection and Research 21: 
45-51. 

Kontschan J. 2010a. Depressorotunda gen. nov., a new 
remarkable Uropodina mite genus from South-East Asia 
with description of four new species (Acari: Mesostigmata). 
Journal of Natural History 44(23-24): 1461-1473. 

Kontschan J. 2010b. Rotundabaloghiid mites of the world 
(Acari: Mesostigmata: Uropodina). Ad Librum Kiado, 
Budapest, 116 pp. 

Kontschan J. 2011. First record of the family Rotundabaloghii- 
dae Hirschmann, 1975 in India, with description of two new 
species of Angulobaloghia Hirschmann, 1975 (Acari: 
Mesostigmata: Uropodina). Opuscula Zoologica (Buda- 
pest) 42(2): 121-124. 

Kontschan J., Stary J. 2011. Uropodina species from Vietnam 
(Acari: Mesostigmata). Zootaxa 2807: 1-28. 

Kontschan J., Stary J. 2012. New Uropodina species and records 
from Malaysia (Acari: Mesostigmata). Acta Zoologica 
Academiae Scientiarum Hungaricae 58(2): 177-192. 


Revue suisse de Zoologie (March 2015) 122(1): 55-70 


ISSN 0035-418 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 


Bernard Landry 


Muséum d'histoire naturelle, C.P. 6434, 1211 Genève 6, Switzerland, Email: bernard.landry@ville-ge.ch 


Abstract: The Pyralidae Pyraustinae of the Galapagos Islands are diagnosed and illustrated and their biology and distri- 
bution are discussed. Of the five species recorded, three are considered as new and described: Neohelvibotys hoecki sp. n., 


Pyrausta galapagensis sp. n., and Pyrausta insolata sp. n. 


Keywords: Achyra - Crambidae - Neohelvibotys - Pyrausta - new species - host plants. 


INTRODUCTION 


The Pyraustinae form the fourth most species-diverse 
subfamily of Pyralidae s. 1.; it is sometimes included in 
family Crambidae (= Crambiformes of Munroe, 1995). 
Pyraustinae are distributed worldwide, and while the 
world fauna includes 190 genera and 1413 species (Re- 
gier ef al., 2012), the Neotropical region is represented 
by 447 species in 35 genera (Munroe, 1995; Nuss er al., 
2014), with Pyrausta being the largest genus with 121 
species in the Neotropical region (Munroe, 1995). The 
Pyraustinae form a strongly supported sister-group rela- 
tionship with the Spilomelinae (Regier et al., 2012), with 
which they were commonly grouped in the past. Diag- 
nostic characters for Pyraustinae are the weakly bilobed 
praecinctorium, atrophied spinula and venulae, male 
forewing with a subcostal retinaculum, male mesothora- 
cic tibia with a hair pencil in a longitudinal groove, male 
tegumen with parallel ridges, male valva almost always 
with a basally directed clasper often accompanied by 
one or more basally or dorsally directed lobes and con- 
spicuous setae or erect scales, and female bursa almost 
always with a rhomboidal or mouth-shaped, spinulose, 
transversely keeled signum (Munroe, 1976; Regier et al., 
2012). 

The Galapagos fauna of Lepidoptera contains close to 
350 species (Roque-Albelo & Landry, 2014) and the 
Pyralidae compose the largest component of the fauna, 
with some 83 species recorded. As with the world fauna, 
the Pyraustinae of the Galapagos occupy the same fourth 
rank in terms of species diversity amongst Pyralidae s. 1., 
albeit with only five species, treated below. 


Manuscript accepted 12.10.2014 
DOI: 10.5281/zenodo.14577 


MATERIAL AND METHODS 


The material investigated here is deposited in the 
American Museum of Natural History, New York, New 
York, USA (AMNH), Natural History Museum, London, 
UK (BMNH), California Academy of Sciences, San 
Francisco, California, USA (CAS), Charles Darwin 
Research Station, Santa Cruz Island, Galapagos, 
Ecuador (CDRS), Carnegie Museum of Natural History, 
Pittsburgh, Pennsylvania, USA (CMNH), where the John 
S. Garth material was deposited, Canadian National 
Collection of Insects, Ottawa, Ontario, Canada (CNC), 
Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts, USA (MCZ), and “Muséum 
d’histoire naturelle de Genève”, Geneva, Switzerland 
(MHNG). The terminology of the structure of the genitalia 
follows Munroe (1976), except for the use of ‘phallus’ 
as recommended by Kristensen (2003). The host plant 
nomenclature follows Jaramillo Diaz & Guézou (2013). 
The manner of giving the label data of the holotypes 
and paratypes is presented in Landry (2006) as are the 
methods used for specimen collecting. 

The previously described species are given additional 
descriptions of the sex and parts that were not treated 
in the original descriptions or elsewhere. The material 
examined for these species is given in alphabetical order 
of islands and within each island as for the paratypes of 
the new species, i.e. in chronological order of date of 
capture. 

The photographs of the moths were taken in the MHNG 
with a Nikon D-800 digital camera and a Nikon Micro- 
Nikkor 105 mm lens. The images were modified in 
Photoshop or Photoshop Elements. Photographs of 
the genitalia were taken with a Leica DFC425 camera 
mounted on a Leica M205C dissecting scope and the 
Leica Application Suite, version 3.7.0. These photos 


56 B. Landry 


were then fused into one with maximal depth of field 
using Zerene Stacker Trial version Build 201305212130, 
with option Align & Stack All (PMax). 


TAXONOMY 
Achyra Guenée, 1849 


Widespread across all major zoogeographical regions 
Achyra includes 18 species (Nuss ef al., 2014), 10 of 
which occur in the Neotropics (Munroe, 1995). The 
larvae feed on a variety of low plants, including some 
crops (alfalfa, beans, clover, cotton, strawberries, etc.) 
(Munroe, 1976). The moths have a conical frons and the 
male genitalia a narrowly triangular uncus, a juxta made 
of two separate plates connected ventrally, a clasper 
with a narrow spinulose process directed ventrally 
at middle or base of sacculus, and the sacculus with a 
blunt process or one or more dorsally directed spines 
at very base and a hump-like process adorned with 
radiating, dorsally directed setae beyond the clasper 
(Munroe, 1976). 


Achyra eneanalis (Schaus, 1923) 
Figs 1-3, 17a-b, 22 

Pyrausta eneanalis Schaus, 1923: 45, 46, fig. 21. — Linsley & 
Usinger (1966: 162). 

Loxostege eneanalis (Schaus). — Capps (1967: 56, 57, figs 53, 
101, 160, 161). 

Achyra eneanalis (Schaus). — Munroe (1976: 46). — Munroe 
(1995: 55). — Roque-Albelo & Landry (2014). — Nuss 
et al. (2014). 


Material examined: 16 £ (4 dissected), 34 © (4 
dissected) from the Galapagos. — Espanola: Bahia 
Manzanillo; Punta Suarez. — Fernandina: Cabo 
Douglas, S 00° 18.269’, W 91° 39.098’; SW side, 
352 m elev{ation]., S 00°20.503’, W 91° 36.969’; Punta 
Espinosa. — Floreana: close to Loberia, 6 m elev., S01° 
17.002’, W 90° 29.460’; close to Las Palmas, 154 m 
elev., S 01° 17.049’, W 90° 28.305’; Punta Cormoran. 
— Gardner near Espanola: NW side. — Genovesa: Bahia 
Darwin. — Isabela: Alcedo, lado NE, low arid zone, 
bosque palo santo; Alcedo, lado NE, 200 m [elev.], 
camp arida alta; Alcedo, NE slope, 292 m elev., S 00° 
23.829’, W 91° 01.957’. — Marchena: [no specific 
locality]. — Pinta: Plaja Ibbeston [sic]; arid zone; + 
15 m elev. — Pinzon: Playa Escondida, 14 m elev., S 
00° 35.928’ W 90° 39.291’. — Plaza Sur: 18 m elev., S 
00° 34.980’, W 09.990’. — Rabida: Tourist trail. — San 


Cristobal: P[uerlto Baquarizo [sic]; near Loberia, sea 
level, S 00°55.277°, W 89° 36.909’; near Loberia, 14 m 
elev., S 00° 55.149’, W 89° 36.897’. — Santa Cruz: 
low agriculture zone, S 00° 42.132’, W 90° 19.156’; 
casa L. Roque-Albelo & V. Cruz, 137 m elev., S 00° 
42.595’, W 90° 19.196’; transition zone, recently cut 
road, S 00° 42.528’, W 90° 18.849’. — Santiago: 
Cerro Inn; La Bomba, 6 m elev., S 00° 11.151’, W 90° 
42.052’. — Seymour Norte: 17 m elev., S 00° 23.935’, 
W 90° 17.369’. Deposited in BMNH, CDRS, CNC, and 
MHNG. 


Diagnosis: In the Galapagos, this medium-sized species 
(14-20 mm wingspan) is unlike any other pyraloid moth 
by virtue of its grey colour and slightly darker faint 
pattern (Figs 1-3) that is sometimes obliterated. One 
unidentified Udea Guenée species (Spilomelinae) in 
the Galapagos is similar in size and sometimes almost 
without apparent pattern on the forewing, but its colour 
is brown and the paler hindwing has a pair of dark 
brown dots at about 1/3 below costa and medially. 


Biology: One specimen from Fernandina (v.1975, 
G. M. Wellington, BMNH) was reared on ‘Cacabus 
miersii’ (Solanaceae). The name of this plant taxon is 
now Exodeconus miersii (Hook. f.) D’Arcy. Widespread 
and endemic in the Galapagos, it is recorded on more 
than 13 islands of the archipelago in the coastal zone 
to the arid lowlands (McMullen, 1999; Jaramillo Diaz 
& Guézou, 2014). Moths have been collected between 
February and May, mostly near sea level, but also up to 
352 m in elevation. 


Distribution: This species is a widespread endemic to 
the Galapagos Islands. The type locality is Conway Bay, 
Santa Cruz Island. Schaus (1923) also reports it from 
South Seymour Island, nowadays more widely known 
as Baltra, and Tower, now Genovesa. I have examined 
or collected specimens from Española, Fernandina, 
Floreana, Genovesa, Isabela, Marchena, Pinta, Pinzon, 
Plaza Sur, Rabida, San Cristobal, Santa Cruz, Santiago, 
and Seymour Norte. 


Remarks: The original description did not take 
genitalia into account and the moth was illustrated 
in black and white. Capps (1967) added details to 
the external morphology description, described and 
illustrated the genitalia, and provided a black and white 
photograph of the moth. The male holotype deposited in 
the USNM (genitalia slide n° 107,347) was studied; the 
male genitalia of my specimens (Figs 17a-b) correspond 
perfectly although there is some apparent variation in 


Figs 1-8. Adults of Galapagos Pyraustinae. (1-3) Achyra eneanalis. (1) 2, Floreana, Punta Cormoran, 21.iv.1992, MHNG. (2) d, 
Isabela, Alcedo, lado NE, 200 m, camp arida alta, 18.iv.2002, MHNG. (3) 9, Fernandina, SW side, 352 m elev., 10.11.2005, 
MUNG. (4-6) Neohelvibotys hoecki. (4) & holotype, CNC. (5) © paratype, Santa Cruz, Tortuga Res., W Sta Rosa, 6.ii.1989, 
CNC. (6) € paratype, Isabela, Alcedo, lado NE, 700 m, camp guayabillos, 16.iv.2002, MHNG. (7, 8) Pyrausta panopealis. 
(7) 3, Santa Cruz, Los Gemelos, 4.v.2002, MHNG. (8) ©, Isabela, 3 km N Sto Tomas, Agr. zone, 8.iii.1989, CNC. 


The Pyraustinae (Lepidoptera, Pyralidae s. |.) of the Galapagos Islands, Ecuador 


57 


58 B. Landry 


the disposition and number of cornuti, but the vesica in 
its devaginated state was not investigated. 


Neohelvibotys Munroe, 1976 


A New World genus containing eight species distributed 
from Maryland, USA, to Argentina (Munroe, 1976, 
1995; Nuss et al, 2014; Patterson ef al, 2014). It 
is recognized in the male genitalia by the narrowly 
triangular uncus, the absence of an oblique ridge 
sporting a row of dorsally directed, often scale-like 
setae medially on the valva, and a single clasper 
(Munroe, 1976). The immature stages are unknown. 


Neohelvibotys hoecki sp. n. 
Figs 4-6, 18a-b, 23 


Material examined: 32 4, 58 ©. 

Holotype: £, ‘ECUADOR [sideways on left side] | 
GALAPAGOS | S[an]ta Cruz, littoral | zone, Tortuga 
Bay | 29.1.1989, M[ercury]V[apour]L[light] | B. Landry 
[/eg.|’; ‘HOLOTYPE | Neohelvibotys | hoecki | B. 
Landry’. Undissected. Deposited in the CNC. 

Paratypes: 31 4, 58 © from the Galapagos Islands, 
Ecuador. — Española: 4 3, 13 ©, Punta Suarez, 10-12. 
i1.[19]67, trap among Lycium minimum, Prosopis doleis 
and Cryptocarpus pyriformis (Gubarsbia) (I. L. Wig- 
gins); 2 € (one dissected, slide MCZ 913), 7 ©, E side of 
peninsula at coast, N of summit, Alt. + 2 m, 18.iv.[19]70 
(R. Silberglied); 3 ©, Bahia Manzanillo, 25.iv.1992, 
M[ercury]V[apour]L[ight] (B. Landry); 2 ©, same data 
except 27.1v.1992; 1 ©, Las Tunas Trail, 100 m elev[a- 
tion]., 30.iv.1992, MVL (B. Landry); 1 ©, Punta Suarez, 
2.v.1992, MVL (B. Landry). — Floreana: 1 3,1 9, Pun- 
ta Cormoran, 21.iv.1992, MVL (B. Landry). — Gardner 
near Espanola: 2 © (one dissected, slide MCZ 914), 
Gardner near Hood, NW Side, Alt[itude]. + 10 m, 17.iv. 
[19]70 (R. Silberglied). — Isabela: 1 £, 11 km N Puerto 
Villamil, 9.111.1989, MVL (B. Landry); 1 4, 1 ©, 8.5 km 
N Puerto Villamil, 11.iii.1989, MVL (B. Landry); 1 d, 
Alcedo, lado NE, 700 m, camp guayabillos, 16.iv.2002, 
uvl (B. Landry, L. Roque); 1 ©, Alcedo, lado NE, 
low arid zone, bosq[ue]. palo santo, 18.iv.2002, uvl 
(B. Landry, L. Roque); 1 © (dissected, slide MHNG 
ENTO 8608), V[olcan]. Darwin, 300 m elev., 15.v.1992, 
MVL (B. Landry); 1 ©, same data except 630 m elev., 
16.v.1992; 1 © (dissected, slide MHNG ENTO 8609), 
same data except 1000 m elev., 18.v.1992. — Pinta: 1 & 
(dissected, slide MHNG ENTO 8611), Plaja [sic] Ib- 
beston [sic], 13.iii.1992, MVL (B. Landry); 1 &, 1 © 
(dissected, slide MHNG ENTO 6590), same data except 
14.11.1992; 1 ©, same data except arid zone, 15.iii.1992; 
1 ©, Cabo Ibbetson, N 00°32.819’, W 90°44.229’, 8 m 
elev., 15.111.2006, uvl (P. Schmitz, L. Roque); 1 ©, 200 m 
elev., 16.111.1992, MVL (B. Landry); 2 ©, same data 
except 400 m elev., 17.iii.1992; 1 4, same data except 


18.11.1992. — Pinzon: 1 3, plaja [sic] Escondida, u[ltra] 
v[iolet]l[ight], 20.iv.2002 (B. Landry, L. Roque). — San 
Cristobal: 2 & (one dissected, slide BL 1801), 2 km SW 
P[uer]to Baquarizo [sic], 11.11.1989, MVL (B. Landry); 
1 3, 4 km SE Pto Baquarizo [sic], 12.11.1989, MVL 
(B. Landry); 1 4, 1 km S El Progreso, 14.ii.1989, 
MVL (B. Landry); 1 4, pampa zone, 15.11.1989, MVL 
(B. Landry); 1 4, Pto Baquarizo [sic], 17.ii.1989, MVL 
(B. Landry); 1 4, 1 ©, base of Cerro Pelado, 22.ii.1989, 
MVL (B. Landry); 1 © (dissected, slide MHNG ENTO 
6589), near Loberia, GPS: elev. 14 m, S 00°55.149’, W 
89°36.897’, 16.i1i.2004, uvl (B. Landry, P. Schmitz); 1 9, 
Chatham Island, 18.iv.[19]32 (M. Willows Jr) (Templeton 
Crocker Expedlition].); 1 9, Puerto Baquerizo Moreno, 
Wreck Bay, v.1975 (T. J. deVries, B.M. 1976-58); 1 9, 
Sappho Cove, Chath[am. Island]. — Santa Cruz: 1 3, Los 
Gemelos, 31.1.1989, MVL (B. Landry); 4 9, C[harles] 
D[arwin]R[esearch]S[tation], arid zone, 3.11.1989, MVL 
(B. Landry); 2 4, 1 ©, Tortuga Res[erve]., W S[an]ta 
Rosa, 6.11.1989, MVL (B. Landry); 1 ©, Media Luna, 
pampa zone, 8.11.1989, MVL (B. Landry); 1 4, Con- 
way Bay, 15.ii.[19]33; 1 4, Media Luna, pampa zone, 
26.11.1989, MVL (B. Landry); 1 &, 2 km W Bella Vista, 
27.11.1989, MVL (B. Landry); 1 9, casa L. Roque-Al- 
belo & V. Cruz, GPS: 137 m elev., S 00°42.595’, W 
090°19.196’, uvl, 27.11.2005 (B. Landry); 2 € (one 
dissected, MHNG ENTO 8610), E[stacion].C[ientifi- 
ca].C[harles].D[arwin]., 4.iii.1992, MVL (B. Landry); 
1 © (dissected, slide BL 1792), Finca S[teve]. Devine, 
17.11.1989, MVL (B. Landry); 1 9, Bahia Conway, 
14.iv.1992, MVL (B. Landry); 1 2, CDRS, Barranco, 
20 m elev., 30.iv.2002, uvl (B. Landry); 1 ¢, Horneman 
Farm, 200 m, 28.vi.1965 (J. DeRoy); 1 © (dissected, BM 
Pyr. Slide n° 21187), Indefatigable [Island], xii.1968 
(B.M. 1969-17, Ref No. L. 44). Deposited in AMNH, 
BMNH, CAS, CDRS, CMNH, CNC, MCZ, and MHNG. 


Diagnosis: In the Galapagos this species is most similar 
to Loxomorpha cambogialis (Guenée), which is also 
often yellow and roughly the same size, but differs 
externally by the conical frons and the hindwing pattern 
without the extra dot submedially. In details of the male 
genitalia this species resembles most closely Loxostege 
subcuprea (Dognin, 1906), stated by Munroe (1995) 
to be misplaced in this genus. However, the wings of 
this species are orange, without pattern except for the 
fuscous costa and apical margin of the forewing and 
most of the fringes of the hindwing (see Capps, 1967: 
43, figs 61, 174). 


Etymology: I take great satisfaction in naming this 
species in honour of Dr Hendrik Hoeck, former director 
of the Charles Darwin Station (1978-1980), founding 
member and president for 17 years, until 2014, of 
the Swiss Friends of the Galapagos Organization, for 
his exceptional dedicated work and devotion to the 
preservation of the fauna and flora of the Galapagos 
archipelago. 


59 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 


Figs 9-16. Adults of Galapagos Pyraustinae. (9-12) Pyrausta galapagensis. (9) ¢ holotype, CNC. (10) £ paratype, Santa Cruz, NNW 
Bella Vista, 225 m elev., 18.ii.2005, MHNG. (11) © paratype, Isabela, NE slope Alcedo, 483 m elev., 31.11.2004, MHNG. 
(12) 3 paratype, same data as Fig. 10, MHNG. (13-16) Pyrausta insolata. (13) 3 holotype, MHNG. (14) € paratype, Isabela, 
NE slope Alcedo, 869 m elev., 2.iv.2004, MHNG. (15) © paratype, Isabela, 3 km N Sto Tomas, Agr. zone, 8.111.1989, CNC. 


(16) © paratype, Pinta, 400 m elev., 17.iii.1992, MHNG. 


60 B. Landry 


Description: MALE (n=32): Head with frons projecting, 
conical, pale burnt orange, with white scales dorsally 
along margin of eye and around ocellus; antennae 
filiform, with flagellomeres slightly thickened, with 
ciliation dense, as long as 1/3 width of flagellomeres, 
dorsal scaling light golden, with white ventrally 
on scape; maxillary palpus burnt orange; labial 
palpus burnt orange dorsally and on whole of third 
segment, white ventrally on first and second segments; 
haustellum white. Thorax dorsally burnt orange at 
collar, progressively paler, yellowish gold, to yellowish 
white. Foreleg coxa, femur and tibia pale golden, 
tarsomeres paler, almost white; tibia slightly swollen, 
with greyish brown hair pencil from medial slit; outer 
spur about half as long as inner spur. Midleg pale 
golden, as on tarsomeres of foreleg, white laterally on 
tibia. Hindleg as midleg, but less evidently white on 
tibia laterally. Forewing length: 7.5-9.0 mm (holotype 
7.5 mm); forewing colour (Figs 4, 6) burnt orange as 
band along costa, otherwise yellowish gold with rest 
of pattern made of slightly darker convex subbasal 
line from slightly before 1/3 costa to almost middle of 
inner margin, postmedian line from 3/4 costa, convex 
to cubital sector, bent straight toward base for short 
distance, then bent at right angle and ending almost 
straight into inner margin, also with short dash at 
apex of cell connecting with costal band, and terminal 
shading; fringe with shorter scales concolorous with 
terminal area of wing, longer scales slightly paler. 
Hindwing pale yellowish gold, with slightly darker 
postmedian band nearly from costa to the first anal 
vein, also with terminal shading; fringe as on forewing. 
Abdomen dorsally pale golden, with white row at apex 
of most segments; ventrally paler, almost white. 

Male genitalia (n=6) (Figs 18a-b). Uncus elongate, about 
as long as tegumen dorsally, flattened, slightly concave, 
gently narrowing from base to narrowly rounded apex, 
with medium-sized setation dorsolaterally from 1/4 ex- 
cept along narrow midline; setae longer near middle 
and changing into thin spines, shortening toward apex. 
Subscaphium lightly sclerotized. Tegumen dorsally with 
more thickly sclerotized, H-shaped (with median bar 
postmedially situated) teguminal ridges projecting api- 
cally on each side of uncus base over connections with 
dorsolateral arms of uncus; laterally with ventral margin 
more thickly sclerotized, forming mediumly-rounded 
apical projection on each side. Valva with dorsal margin 
subbasally humped, after connection with transtilla, then 
straight; with medially bare, more thickly sclerotized 
band along costa narrowing to 2/3, with a few long se- 
tae and scales laterally; ventral margin straight to 1/10, 
then angled and shortly straight again, then only slightly 
projecting to broadly rounded apex, with few long se- 
tae especially toward base; clasper with ventral process 
simple, claw-like, at 1/3 length of valva, accompanied by 
short, more or less narrowly rounded basal lobe pointing 
toward costa of valva and set with short to long setae; 


sacculus a low, broadly-rounded hump set with short se- 
tae. Transtilla thin, unsclerotized medially. Diaphragma 
ventrad from transtilla with patch of about 60 short setae 
on each side; with one separate seta closer to middle on 
each side at level of base of setose patches. Juxta short, 
slightly wider than long, with broadly rounded basal 
margin, laterally extending into short, thin projections, 
with apical margin broadly concave. Vinculum narrow, 
with short saccus. Phallus straight, slightly longer than 
tegumen + uncus, more thickly sclerotized on left side 
on basal half, with narrow sclerotized band from mid- 
dle on left side running in diagonal dorsally toward apex, 
enlarging to 2 X wider, twisting sheath subapically and 
ventrally, ending in long, narrow hook curving upward 
laterally on right side and reaching dorsal margin of 
shaft; ventrally at apex with short, down-curbed rounded 
‘lip’; coecum penis short, rounded; vesica with elongate 
(about 1/4 length of shaft) band of 35-40 thick and short 
cornuti, and smaller, more rounded patch of more or less 
conspicuous smaller cornuti situated slightly more basad 
in invaginated condition; ductus ejaculatorius forming 
about 12 coils. 

FEMALE (n=58) (Fig. 5): Antenna thinner and less 
densely ciliate than male’s. Forewing length: 7.0-9.0 mm. 
Female genitalia (n=5) (Fig. 23). Papillae anales narrow, 
curved (apically projecting dorsally and ventrally), 
abundantly setose on whole surface, with laterobasal 
sclerotized narrow band about twice as wide ventrally, 
not connecting either dorsally or ventrally; posterior 
apophyses thin, pointed, with short basal projection 
ventrally, about 2/3 length of papillae anales. Segment 
VIII of medium length, 4-5 X length of sclerotized band of 
papillae anales at level of apophyses, ventrally reducing in 
width, forming elongate triangle, not connected medially, 
with 6-8 short setae on each side dorsally; anterior 
apophyses about 1.6 X length of posterior apophyses, 
with short, narrowly triangular ventral projection at 1/4th 
of length from base. Lamella postvaginalis covered with 
spinules except along narrow midline, forming wide 
triangle across width of segment with cephalad margin 
slightly bent posterad; laterally and anteriorly from 
subbasal ventral projections of anterior apophyses with 
sclerotized band of medium width spinulose posteriorly. 
Antrum funnel-shaped, lightly sclerotized laterally, 
with sclerotized ridges extending into ductus bursae. 
Basal section of ductus bursae of medium girth, dorsally 
sporting flat, accessory sac about 1.5 X as long as width 
of ductus at its level, with sclerotized ridges ending at 
more heavily sclerotized, dorsally open collar of about 
1/8 length of corpus bursae; subsequent section of corpus 
bursae a more or less triangular and thickly sclerotized 
widening which connects with ductus seminalis; final 
section of ductus bursae made of 6 coils of medium 
girth. Corpus bursae circular, not very large, apically (at 
bottom) with more or less thickly sclerotized small to 
medium-sized rounded signum with scobinations. 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 61 


Biology: Unknown except for the nocturnal habits of 
the moths, which are attracted to light, and the diversity 
of habitats in which they fly, from sea level to 1000 m in 
elevation. 


Distribution: Galapagos islands of Española, Floreana, 
Gardner near Espafiola, Isabela, Pinta, Pinzon, San 
Cristobal, and Santa Cruz. 


Remarks: The species keys out as a member of 
Neohelvibotys in Munroe (1976). However, the key 
provided by Munroe mostly uses genitalia characters of 
the males, and this species does not entirely agree with 
the diagnosis (no formal description was given) of the 
genus with respect to the female genitalia. According 
to the diagnosis, the ostial chamber is large and the 
ductus seminalis does not form a broad arch with the 
ductus bursae as in Achyra and Hahncappsia. Also, the 
species of Neohelvibotys illustrated by Capps (1967) 
and Munroe (1976) have a small appendix bursae, 
absent in N. hoecki, while this species has an appendix 
(accessory sac) of the ductus seminalis, absent in 
the other species. Possibly this species is aberrant in 
female genitalia characters, when compared with the 
type species [N. neohelvialis (Capps)] and the other 
Neohelvibotys species for which the female genitalia 
are known, or the genera are yet improperly defined 
and need a phylogenetic analysis. I have checked all 
described species of Achyra Guenée, Hahncappsia 
Munroe, Helvibotys Munroe, Neohelvibotys Munroe, 
and species stated to be misplaced in Loxostege Hübner 
recorded from the Neotropical region by Munroe (1995) 
to make sure that this species was not already described. 
The dissected male and female MCZ paratypes from 
Espafiola were dissected by Eugene G. Munroe in 
October 1983. The labels on the slide preparation of 
these specimens identify them as Neohelvibotys in 
Munroe’s hand-writing. 


Key to the Galapagos species of Pyrausta Schrank 


Pyrausta Schrank, 1802 


A widely distributed genus comprising 284 species 
(Nuss er al, 2014). The larvae web and often roll 
leaves, shoots, and flowers head of a variety of 
herbaceous plants, but most of the species were reared 
from Lamiaceae (Munroe, 1976). Three species occur 
in the Galapagos. The moths are generally small and 
slender and the valva of the male genitalia usually bears 
a single basally directed or recurved setose or scaled 
clasper (Munroe, 1976). 


Pyrausta panopealis (Walker, 1859) 
Figs 7, 8, 19a-b, 24 


Rhodaria panopealis Walker, 1859: 318. 

Pyrausta panopealis Walker. — Munroe (1976: 111, 112, not il- 
lustrated). —Munroe (1983: 71) — Robinson et al. (1994: 
179, pl. 29 fig. 19). — Shaffer et al. (1996: 189). — Peck 
et al. (1998: 227). — Munroe (1995: 57). — Heppner 
(2003: 275). — Causton er al. (2006: 141). — Roque-Al- 
belo & Landry (2012). — Nuss et al. (2014). — Patterson 
et al. (2014). 


Material examined: 14 © (4 dissected), 16 © 
(3 dissected). — /sabela: 3 km N S[an]to Tomas, 
Agriculture zone; 11 km N Pfuer]to Villamil; NE 
slope Alcedo, near shore, GPS: elev[ation]. 9 m, S 
00°23.619’, W 90° 59.715’; NE slope Alcedo, GPS: 
elev. 292 m, S 00° 23.829’, W 91° 01.957’; Alcedo, lado 
NE, camp arida alta, 200 m [elev.]; V[olcan]. Darwin, 
300 m elev. — Santa Cruz: 4 km N Puerto Ayora; casa 
L. Roque-Albelo & V. Cruz, GPS: 137 m elev., S 00° 
42.595’, W 90° 19.196’; transition zone, recently cut 
road, GPS: S 00° 42.528’, W 90° 18.849”, [reared] from 
Hyptis sidaefolia leaves; low agriculture zone, GPS: 00° 
42.132’, W 90° 19.156’; Finca Vilema, 2 km W Bella 
Vista; Los Gemelos. — Santiago: N side, GPS: 437 m 
elev., S 00° 13.316’, W 90° 43.808’; Cerro Inn; Bahia 


Il? 


DO 


Forewing with orange spots or mostly orange; if forewing pattern obscured, then hindwing paler base with orange 
scaling; male genitalia with uncus of medium girth, with or without long setae along ventral margin of valva at 
base; female genitalia with colliculum of ductus bursae followed by distinctly enlarged section .......................... 2 
Forewing usually with mellow yellow markings; if forewing pattern obscured or with orange scaling, then hind- 
wing paler base light yellow (Figs 9-12); male genitalia with narrow uncus and very long setae along ventral 
margin of valva at base; female genitalia with colliculum of ductus bursae followed by almost equally narrow 
SECLO PM RE N RR EEE P galapagensis sp. n. 
Forewing darker colour violet, with subapical orange marking extending along costa to apex; orange markings 
usually forming 4 spots, the median two sometimes converging into one; forewing base orange instead of darker 
violet, before postbasal line (Figs 7, 8); male genitalia with clasper distinctly spoon shaped; corpus bursae of fe- 
malezeenitaltaswithfappendixaburs a Ce P. panopealis (Walker) 
Forewing darker colour dark brown, with subapical orange marking not extending to apex along costa; orange 
more extensive, especially medially and at base, sometimes with dark brown reduced to lines (postbasal, postme- 
dian, and subapical) and dash at end of cell, except in less common specimens with forewing entirely dark (Figs 
13-16); male genitalia with clasper a simple, short, triangular sheet; corpus bursae of female genitalia without 
APPENAISGDUTSACHS EEE cere ee III I TI Rivara P insolata sp. n. 


62 B. Landry 


Espumilla; 200 m elev.; Aguacate, 520 m elev.; Central, 
700 m elev.; Jaboncillo, + 850 m elev. Deposited in 
CDRS, CNC, MHNG. 


Diagnosis: In the Galapagos this small species can only 
be confused with Pyrausta insolata sp. n., but the key 
above will separate specimens of these two. Outside 
of Galapagos the species is most similar to Pyrausta 
phoenicealis (Hübner), but on average smaller (6-7 mm 
in forewing length), with wings lighter, more thinly 
scaled, the forewing with dark areas redder and light 
areas yellower, the hindwing lighter and not as red as 
the forewing, with the pale areas weakly contrasting 
and without a fulvous patch at base, the fringes with 
the basal parts yellow or with a few reddish scales and 
distal parts whitish, etc. (Munroe, 1976). The female 
hindwing is generally darker than that of the male 
because the lighter orange markings are smaller than 
those of the male. 


Biology: Hyptis capitata Jacquin (Lamiaceae) was 
reported as a host plant in Puerto Rico (Munroe, 1976) 
and Dicerandra frutescens Shinners and Hyptis capitata 
Jacquin (Lamiaceae) in Florida (Heppner, 2003). I have 
reared specimens from Hyptis sidaefolia (L’Hér.) Brig. 
(Lamiaceae) leaves on Santa Cruz Island. The moths are 
attracted to light and have been collected between the 
coastal zone and up to 850 m in elevation. 


Distribution: Described from China, this species 
is widespread in the tropics of the world (Munroe, 
1976). In the Galapagos I have collected specimens 
on the islands of Isabela, Santa Cruz, and Santiago. 
One specimen in the CDRS with a label stating ‘41’ 
was associated with catalogue data stating ‘Pyrausta 
phoenicealis, Santa Cruz, Fernandina, Isabela, Santiago, 
Transition to humid zones, Feb[ruary]., June, July, 
Dec[ember].’ 


Remarks: For the full synonymy pertaining to 
this taxon see Munroe (1976, or 1995). I confirm 
that the holotype deposited in the BMNH is a 
female. Pyrausta panopealis has been confused and 
synonymized with Pyrausta phoenicealis (Hübner, 
1818) in some publications prior to Munroe (1976), 
who removed it from synonymy based on small 
differences (see Diagnosis). Later as well (see for 
example Guillermet, 2009, and Vari et al., 2002), the 
species has been reported as P phoenicealis, with 
P. panopealis in synonymy, although Munroe (1976) 
stated that P phoenicealis ‘seems to be confined to the 
southeastern part of the United States [of America]’ 
and its type locality is Florida. I have decided to 
follow Munroe (1976, 1983, 1995) in considering that 
P. panopealis and P phoenicealis are different taxa 
as he made the only available comparative study of 
the group and because there are DNA COI barcode 
unpublished data that seem to support the separation of 
the two species (J.-F. Landry, pers. comm.). The host 


plant record of Hyptis capitata Jacquin (Lamiaceae) by 
Munroe (1976) is based on Schaus (1940) who reported 
the species as P phoenicealis from Puerto Rico. 


Pyrausta galapagensis sp. n. 
Figs 9-12, 20a-b, 25 


Material examined: 7 4, 15 ©. 

Holotype: ©, ‘ECUADOR [sideways on left side] | 
GALAPAGOS | Isabela, 3 km N. | S[an]to Tomas, Agr|i- 
culture] Zone | 8.III.1989, M[ercury]V[apour]L[light] | 
B. Landry [/eg.]|’; ‘HOLOTYPE | Pyrausta | galapagensis 
| B. Landry’. Undissected. Deposited in the CNC. 
Paratypes: 6 4, 15 © from the Galapagos Island, Ecua- 
dor. — Floreana: 1 & (dissected, slide B.M. Pyralidae 
21185), 1 ©, Charles [Island], Asilo de la Paz, 360 m, 
i.1971, BM 1971-79, Ref. No. L-159. — Isabela: 1 & (dis- 
sected, slide PYR 374), 1 ©, same data as holotype; 1 9, 
NE slope Alcedo, near pega-pega camp, GPS: elev[a- 
tion]. 483 m, S 00°24.029’, W 91°02.895’, 31.11.2004, 
ulltra]v[iolet]l[fight] (B. Landry, P. Schmitz); 1 © (slide 
MHNG ENTO 8622), NE slope Alcedo, ca. 400 m up (S) 
Los Guayabillos camp, GPS: elev. 892 m, S 00°24.029’, 
W91°04.765’, 1.iv.2004, uvl (B. Landry, P. Schmitz); 
1 4, 1 ©, Alcedo, lado NE, 400 m, camp pega-pega, 
15.iv.2002, uvl (B. Landry, L. Roque). — Santa Cruz: 
2 ©, Media Luna, pampa zone, 21.1.1989, M[ercury] 
V[apour]L[ight] (B. Landry); 1 ©, casa L. Roque-Al- 
belo & V. Cruz, GPS: 137 m elev., S 00°42.595’, W 
090°19.196’, 17.11.2005, day time (B. Landry); 2 € (one 
dissected, slide MHNG ENTO 8620), 4 © (one dissec- 
ted, slide MHNG ENTO 8621), NNW Bella Vista, GPS: 
225 m elev., S 00°41.293’, W 090° 19.665’, 18.11.2005, 
uvl (B. Landry, P. Schmitz); 1 ©, Los Gemelos, 4.v.2002, 
uvl (B. Landry, L. Roque); 1 4, 1 ©, Indefatigable [Is- 
land], v.1970 (R. Perry & Tj. De Vries), B.M. 1970-371, 
Ref. No. L. 150; 1 9, same data except ‘B.M. 1969-17’. 
Deposited in BMNH, CDRS, CNC, and MHNG. 


Diagnosis: In forewing pattern and colour the 
paler specimens of this species are most similar to 
P. deidamialis (Druce), the lectotype of which is from 
Volcan Chiriqui, Panama. In male genitalia this new 
species differs from P deidamialis in the distinctly 
narrower valvae and uncus, the spatulate clasper, and 
the three cornuti of the vesica. The female genitalia of 
P. deidamialis are unknown to me. In the Galapagos this 
species stands out from the other two Pyrausta species 
present, which have conspicuous orange scaling (see 
Key above). 


Etymology: The name refers to the area of distribution, 
the Galapagos Archipelago. 


Description: MALE (n=7): Head with frons slightly 
projecting, apically flat, burnt orange in middle, dark 
brown along the sides, with white band laterally from 
base of antennae to above frons and also ventral from 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 63 


Figs 17-19. Male genitalia of Galapagos Pyraustinae, a and b always from same slide, but not to scale, a showing the genitalia without 
phallus and b, the phallus. (17) Achyra eneanalis: slide MHNG ENTO 8616, Fernandina. (18) Neohelvibotys hoecki 
paratype: slide MHNG ENTO 8610, Santa Cruz. (19) Pyrausta panopealis; slide CNC PYR 373, Santa Cruz. bss: basal 
spine of sacculus; c: clasper; hps: hump-like process of sacculus. 


64 B. Landry 


compound eye; antennae filiform, with ciliation dense, 
as long as 1/3 width of flagellomeres, with one single 
longer (half as long as corresponding flagellomere) seta 
dorsally on about first 10 flagellomeres, scaling mostly 
greyish brown, with some white, dark brown and burnt 
orange on scape and pedicel; maxillary palpus burnt 
orange or dark brown with few burnt orange scales; 
labial palpus burnt orange, sometimes with dark brown, 
on most of second segment and all of third, satiny 
white ventrally on second segment and all of first; 
haustellum light yellow to light orange and sometimes 
greyish brown. Thorax dorsally at collar burnt orange, 
sometimes with longitudinal darker lines on scales, to 
lighter royal yellow, progressively becoming lighter 
toward apex, sometimes with brown scales, dirty white 
to light yellow on thoracic segment III. Foreleg coxa 
light greyish brown with few burnt orange and dirty 
white scales; femur burnt orange; tibia dark greyish 
brown with white laterally and some burnt orange; 
tarsomeres light greyish brown, with some white at 
base of first tarsomere, progressively becoming darker 
greyish brown. Midleg coxa satiny white; femur light 
gold, with burnt orange and dark brown at tip; tibia 
dorsally at base with dark brown and burnt orange, 
progressively becoming paler toward apex, laterally 
satiny white; tarsus light gold with greyish white on 
last two tarsomeres. Hindleg coxa white, otherwise 
leg entirely light gold. Forewing length: 6.0-7.0 mm 
(holotype 6.0 mm); forewing colour (Figs 9, 12) dark 
brown on costa and second half of wing, more or less 
speckled with burnt orange, base of wing more mellow 
to light yellow toward dorsum; dark brown markings 
as submedian line straight from dorsum to base of cell 
where one or two more or less distinct dark brown spots 
may be present, postmedian line from dorsum straight to 
middle of wing where it curves externally to sometimes 
reach broad terminal band, and short bar at end of cell; 
mellow yellow markings as pair of spots or complete 
line from dorsum, but not reaching costa, bordering 
postmedian dark line, and in cell and sometimes below 
until dorsum; fringe with shorter basal scales dark 
greyish brown and longer scales lighter greyish brown. 
Hindwing base light yellow, often with dark brown spot 
submedially in median sector and straight line from 
median sector to anal sector postmedially, followed 
by light yellow before broad apical dark brown band; 
fringe as on forewing. Abdomen dorsally mostly light 
greyish brown speckled with light yellow or orange and 
with white line at apices of segments II-VII, although 
last two segments mostly light gold; ventrally light gold 
with some light greyish brown on sternites V-VII. 

Male genitalia (n=3) (Figs 20a-b). Uncus elongate, 
about 1.6 X length of tegumen dorsally, flattened, very 
slightly down-turned, narrowing from base to about 
2/5, then parallel-sided until slightly pointed apex, with 
medium-length setation laterally from about middle to 
3/4, with short, thick setae from beyond middle dorsal- 


ly until apex. Subscaphium lightly sclerotized, shorter 
than uncus. Tegumen short, dorsally with more thickly 
sclerotized teguminal ridges diverging apically and ba- 
sally, without distinct median bar connecting them, but 
rather with scale-less area slightly shorter than tegumi- 
nal ridges; laterally with more thickly sclerotized ven- 
tral margin slightly concave, with scaled area between 
it and teguminal ridges rounded, more thickly scaled to- 
ward dorsoapical margin. Valva narrow, with dorsal mar- 
gin slightly humped at connection with transtilla, then 
slightly concave; medially bare and thickly sclerotized 
band along costa narrowing to 2/3, with some moder- 
ately long thin setae and scales on dorsal edge; ventral 
margin straight at base, subbasally angled, then straight 
until rounded apex; medially with few short thin scales 
at base; laterally, next to ventral margin, and along ven- 
tral margin with about 10 moderately long to very long 
setae between 1/10 to 1/3; clasper directed basomedially, 
shaped like racquetball racquet, with mostly short and 
thick, apically unmodified setae, but also a few longer 
ones mostly along apical and lateral margins; sclerotized 
plate before clasper medially forming broad, low triangle 
with short, narrow, rounded point set with about 10 short 
setae; sacculus unmodified. Transtilla thin, unsclerotized 
medially. Diaphragma ventrad from transtilla with patch 
of about 10 medium-length setae on each side. Juxta 
short, quadrangular, almost completely flat, with low 
depression medially, with apical margin projecting and 
medially with short v-shaped cut, with basal margin at 
each angle with short, rounded projection. Vinculum nar- 
row with short saccus apically narrowly rounded. Phallus 
an almost straight tube, about 10% longer than tegumen 
+ uncus, basal half thickly sclerotized only ventrally; 
coecum penis absent; vesica with three cornuti: longest 
half as long as phallus shaft, second 3/5 length of longest, 
third 2/5 length of longest; ductus ejaculatorius forming 
one coil distally. 

FEMALE (n=15) (Figs 10, 11): Antenna thinner than that 
of the male. Forewing length: 6.5-7.5 mm. 

Female genitalia (n=2) (Fig. 25). Papillae anales short, 
slightly convex, moderately setose, with short to mode- 
rately long setae mostly along basal (in one line) and apical 
margins; cuticle covered with spinules, with laterobasal 
sclerotized band narrow, wider at bases of apophyses, 
not connecting either ventrally or dorsally; posterior 
apophyses long and thin, without ventral projection at 
base, about 1.5 X width of papillae anales. Segment VIII 
short, 2 (ventrally) — 3 (dorsally) X length of sclerotized 
band of papillae anales at level of apophyses, ventrally 
not connected in middle, with few short to medium-length 
setae mostly along apical margin and dorsally, covered 
with spinules; anterior apophyses about 0.85X length of 
posterior apophyses, with dorsal, triangular enlargement 
at about 1/4. Membrane unspecialized around ostium 
bursae. Ductus bursae with membranous and spinulose 
funnel-shaped antrum about 1/5 of total length, followed 
by shorter colliculum devoid of spinules, followed by 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 65 


short membranous lateral extension and constriction 
abundantly adorned with thick spinules, followed by 
sclerotized and ridged wider section devoid of spinules 
and about 1/3 length of whole structure, with lateral 
rounded projection at base, followed by membranous, 
narrower, ridged and bent section devoid of spinules and 
slightly enlarging into corpus bursae. Ductus seminalis 
connecting between second sclerotized section of 
ductus bursae and next. Corpus bursae circular, with 
spinules either single or in rows of 2-6 except distad of 
signum; signum very large, shaped like closed pair of 
thick, stylized human lips, with two low crests directed 
internally on each side of middle along midline, which 
thus appears more thickly sclerotized, with perpendicular 
scale-like extensions from middle, and with thick, short 
spines directed internally; with accessory sac about half 
of size of corpus bursae, devoid of spinules. 


21b 


Biology: Unknown except that moths are attracted 
to light and that the species seems to prefer habitats 
situated at medium (225 m) to high elevations, up to the 
pampa zone. 


Distribution: Galapagos Islands of Floreana, Isabela, 
and Santa Cruz; probably endemic. 


Remarks: The females are often darker than males. The 
colour definitions are partly from Wikipedia. 


Pyrausta insolata sp. n. 
Figs 13-16, 21a-b, 26 


Material examined: 26 4, 21 ©. 

Holotype: ©, ‘ECU[ADOR]., GALAPAGOS | Pinta, 
17.11.1992 | 400 m elev[ation]., M[ercury]V|apour] 
L[ight] | /eg[it]. B. Landry’; “HOLOTYPE | Pyrausta | 


Figs 20-21. Male genitalia of Galapagos Pyrausta species paratypes, a and b from same slide, but not to scale, a showing the genitalia 
without phallus and b, the phallus. (20) P galapagensis: slide CNC PYR 374, Isabela. (21) P insolata: slide MHNG 
ENTO 8628, Santiago. 


66 B. Landry 


insolata | B. Landry’. Undissected. Deposited in the 
MHNG. 

Paratypes: 25 ¢, 21 © from the Galapagos Island, 
Ecuador. — Fernandina: 1 & (dissected, slide MHNG 
ENTO 8631), SW side, GPS: 815 m elev{ation|. 
S 00° 21.270’, W 091° 35.341’, 11.11.2005, u[ltra]v[io- 
let]l[ight] (B. Landry, P. Schmitz); 1 4, SW side, crater 
rim, GPS: 1341 m elev., S 00° 21.910’, W 091° 34.034’, 
12.11.2005, uvl (B. Landry, P. Schmitz); 1 4, same data 
except 13.11.2005; 1 4, [no precise locality], iii.1970 
(Perry & de Vries). — Isabela: 3 © (one dissected, slide 
PYR 372), 3 km N S[an]to Tomas, Agrliculture]. Zone, 
8.iii. 1989, M[ercury]V[apour]L[ight] (B. Landry); 2 © 
(one dissected, slide PYR 353), 1 ©, 11 km N Puerto Vil- 
lamil, 9.iii.1989, MVL (B. Landry); 3 © (one dissected, 
slide MHNG ENTO 8632), 1 ©, NE slope Alcedo, ca. 
400 m up (S) Los Guayabillos camp, GPS: 892 m elev., 
S 00° 25.208’, W 91° 04.765’, 1.1v.2004, uvl (B. Landry, 
P. Schmitz); 1 4, 1 9, NE slope Alcedo, Los Guayabillos 
camp, GPS: 869 m elev., S 00° 24.976’, W 91° 04.617’, 
2.1v.2004, uvl, 4h00-5h30 (B. Landry, P. Schmitz); 
1 4, 1 ©, Alcedo, lado NE [side], 400 m [elev.], camp 
pega-pega, 15.iv.2002, uvl (B. Landry, L. Roque); 1 4, 
1 ©, Alcedo, lado NE, 700 m [elev.], camp guayabil- 
los, 16.iv.2002, uvl (B. Landry, L. Roque); 1 ©, Alce- 
do, lado NE, 1100 m [elev.], cumbre [summit], caseta 
Cayot [cabin], 17.1v.2002, uvl (B. Landry, L. Roque); 
3 ©, V[olcan]. Darwin, 630 m elev., 17.v.1992, MVL 
(B. Landry); 1 3, + 15 km N Pto Villamil, 25.v.1992, 
MVL (B. Landry); 2 ©, Sierra Negra, Corazon Verde, 
xi-xii.1974 (T.J. De Vries, B.M. 1976-58). — Pinta: 1 3, 
372 m elev., 16.11.2006, N 00° 34.476’, W 90° 45.102’, 
uvl (P. Schmitz, L. Roque); 1 ¢, 1 © (dissected, slide 
MHNG ENTO 8629), 200 m elev., 16.11.1992, MVL 
(B. Landry); 2 4,3 ©, same data as holotype; 1 4,400 m 
elev., 18.iii.1992, MVL (B. Landry); 2 © (one dissec- 
ted, slide BL 1248), 400-650 m elev., day (B. Landry). 
— Santa Cruz: 2 ©, Tortuga Res[erve]. W S[an]ta Rosa, 
6.11.1989, MVL (B. Landry). — Santiago: 1 3, N side, 
GPS: 437 m elev., S 00° 13.316’, W 90° 43.808’, 
3.111.2005, uvl (P. Schmitz); 1 © (dissected, slide MHNG 
ENTO 8628), 1 © (dissected, slide MHNG ENTO 8630), 
NE side, close to caseta, GPS: 686 m elev., S 00°14.177’, 
W 90°44.619°, 6.111.2005, uvl (P. Schmitz); 1 4, 1 © (dis- 
sected, B.M. Pyralidae slide 21186), (James), vii.1970 
(B.M. 1970-567, Ref. No. L 159); 1 ©, (James), Bahia 
Bucanero, iv.1974 (B.M. 1975-7, Ref. No. L [blank]). 
Deposited in BMNH, CDRS, CNC, and MHNG. 


Diagnosis: This species is most similar to Pyrausta 
insignitalis (Guenée), P onythesalis (Walker) and 
P. pseudonythesalis Munroe, but it resembles the former 
more in habitus by virtue of the predominance of orange 
colouration. In male genitalia (see Munroe, 1976, pl. J 
fig. 6 for a photo of the male genitalia of P insignitalis), 
the new species differs in the shorter and broader 
uncus, absence of a thin projection of the clasper, 
and sclerotized plate before clasper without dorsal 


projection. In female genitalia the new species differs 
especially in the wide and singly coiled ductus bursae. 
In the Galapagos this species is most similar to the other 
two Pyrausta species recorded and the key presented 
above will separate the three. 


Etymology: From Latin, meaning ‘exposed to the sun’; 
in reference to the colour of the moths and the fact that 
the species being present on the equatorial Galapagos 
archipelago indeed receives a lot of sunshine. 


Description: MALE (n=26): Head with frons slightly 
projecting, apically slightly rounded (not quite flat), 
orange except for few whitish yellow scales at base 
of antenna; antennae filiform with ciliation dense, 
as long as 2/3 width of flagellomeres, with one single 
longer (2/3 as long as corresponding flagellomere) 
seta dorsally on about first 10 flagellomeres, scaling 
orange on scape and pedicel, greyish brown to whitish 
grey on flagellum; maxillary palpus greyish orange; 
labial palpus orange, with paler whitish yellow scales 
ventrally on first segment and burnt orange dorsally on 
second segment and apically on third; haustellum light 
orange. Thorax dorsally orange to light burnt orange 
at collar and bases of patagia, sometimes with light 
brown, orange in middle and slightly lighter orange 
on thoracic segment III. Foreleg coxa light golden to 
light orange; femur as coxa, with orange at tip; tibia 
greyish brown with light golden laterally and at tip; 
tarsomeres light golden and greyish brown dorsally. 
Midleg coxa whitish golden; femur light golden with 
orange at tip; tibia orange dorsally with light greyish 
brown at base and dorsally on spurs, light golden 
elsewhere; tarsomeres light golden with light greyish 
brown at base of first tarsomere. Hindleg light golden 
with light orange on tibia dorsally and light greyish 
brown on tarsomeres dorsally. Forewing length: 6.0- 
7.0 mm (holotype 7.0 mm); forewing colour (Figs 
13, 14) mostly orange, with dark brown on costa as 
fine line, complete submedian slightly convex line, 
sometimes one small spot at base of cell, one distinct 
straight bar at end of cell usually associated with more 
or less intense suffusion between bar and terminally 
bulging section of postmedian line, small triangle on 
costa at postmedian line, straight subterminal line often 
with small indentations at CuA, and M,, often fused 
with terminal brownish black spots at apices of veins 
to form wider terminal band, sometimes whole wing 
with heavier suffusion and broad costal band (Figs 15, 
16); fringe with basal scales forming pale orange and 
greyish brown spots alternately or mostly dark greyish 
brown in darker specimens, longer scales pale greyish 
brown. Hindwing orange with dark brown markings as 
small spot in cell, postmedian line, subterminal band 
often fused with terminal band, and suffusion usually 
between CuP and 2A up to middle of wing; fringe 
similar to that of forewing. Abdomen dorsally uniformly 
orange or with greyish brown on segments I-V in darker 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 67 


Figs 22-26. Female genitalia of Galapagos Pyraustinae. (22) Achyra eneanalis: slide MHNG ENTO 8617, Fernandina. (23) 
Neohelvibotys hoecki paratype: slide BL 1792, Santa Cruz, CNC. (24) Pyrausta panopealis: slide MHNG ENTO 8624, 
Santiago. (25) P galapagensis paratype: slide MHNG ENTO 8621, Santa Cruz. (26) P insolata: a- slide CNC PYR 372, 
Isabela; b- signum, slide BM Pyralidae n° 21186, Santiago. ab: appendix bursae; as: accessory sac; co: colliculum; odb: 
outgrowth of ductus bursae. 


68 B. Landry 


specimens, ventrally light golden, with light orange on 
distal segment in darker specimens. 

Male genitalia (n=5) (Figs 21a-b). Uncus stout, about 
1.6 X length of tegumen dorsally, flattened and straight 
in lateral view, with dorsal margin apically down-turned, 
with lateral margins gently converging to mediumly 
rounded apex, with 1-2 long setae dorsolaterally near 
middle, apical third covered with medium-length to 
short thickened setae decreasing in size towards apex. 
Subscaphium with narrow, lightly sclerotized ventral 
margin reaching apex of uncus. Tegumen short, dorsally 
with more thickly sclerotized teguminal ridges diverging 
apically and basally, H-shaped, with more thickly 
sclerotized horizontal bar present, forming basal margin, 
with scale-less area between teguminal ridges 3/5 length 
of latter; laterally with more thickly sclerotized ventral 
margin slightly concave, with scaled area between it 
and teguminal ridges square, more thickly scaled toward 
dorsoapical margin. Valva of medium width, with dorsal 
margin slightly humped at connection with transtilla, 
then slightly concave; medially bare and_ thickly 
sclerotized band along costa narrowing to 1/3 of length 
of valva, then parallel-sided until ending at 3/4 length of 
valva, with some moderately long thin setae and scales 
on dorsal edge; ventral margin not distinctly straight at 
base, subbasally broadly rounded, then straight to mid- 
length and slightly convex to rounded apex; medially 
with few short thin scales at base; laterally, next to 
ventral margin, and all along ventral margin with short 
to long setae; clasper a simple, short, triangular sheet 
with dorsal margin produced and rounded, medially 
covered with short to long thickened setae decreasing 
in length toward apex; sclerotized plate before clasper 
broadly rounded, not projected dorsally beyond dorsal 
angle of clasper, adorned with 8-10 moderately long 
setae; sacculus unmodified. Transtilla thin, unsclerotized 
medially. Diaphragma ventrad from transtilla with patch 
of about 6-10 medium-length setae on each side. Juxta 
short, shaped like inverted human pelvis. Vinculum 
narrow, with short, laterally compressed saccus with 
blunt apical margin. Phallus a slightly curved, short tube 
about 20% longer than tegumen + uncus, basal 1/3 thickly 
sclerotized only ventrally; coecum penis short, rounded; 
vesica with a long, thin, curved cornutus about 3/5 length 
of phallus, one thicker, stiletto-like cornutus about half as 
long, and about 5 additional thin cornuti also about half 
as long; ductus ejaculatorius forming one coil. 

FEMALE (n=21) (Figs 15, 16): Antenna thinner than that 
of the male. Wings usually with more dark brown scaling 
than those of males, sometimes with dark brown and also 
sometimes burnt orange scaling completely obscuring 
orange areas. Forewing length: 6.0-7.0 mm. 

Female genitalia (n=3) (Fig. 26). Papillae anales 
short, slightly convex, abundantly setose, with short 
to moderately long setae, the latter mostly along basal 
margin; cuticle covered with spinules, with laterobasal 
sclerotized band narrow, wider near bases of posterior 


apophyses, not connecting either ventrally or dorsally; 
posterior apophyses short, slightly thicker at base, about 
as long as width of papillae anales. Segment VIII short, 
with sclerotized section a narrow band of nearly equal 
length ventrally and dorsally, 2 X length of sclerotized 
band of papillae anales at level of apophyses, ventrally 
not connected in middle, with few medium-length 
setae mostly along apical margin laterally and dorsally, 
covered with spinules; anterior apophyses almost 2 X 
length of posterior apophyses, with dorsal, triangular en- 
largement at about 2/5. Membrane unspecialized around 
ostium bursae, covered with spinules. Ductus bursae 
with short, membranous and spinulose funnel-shaped 
antrum, followed by equally short sclerotized incomplete 
(open dorsally) colliculum devoid of spinules, followed 
by enlarging membranous section forming one twist, 
4 X as long as first two sections, almost 4 X as wide 
as colliculum, adorned with elongate sclerotized and 
wrinkled outgrowth decreasing in size from base to 
middle, lightly spinulose. Ductus seminalis connecting 
between second, sclerotized section of ductus bursae and 
next. Corpus bursae short, circular, not much wider than 
ductus bursae, about half as long as ductus bursae, lightly 
spinulose; signum a small, square plate with short spines 
projecting inside, with or without depression in middle; 
without accessory sac. 


Biology: Unknown except that moths are attracted 
to light and that the species seems to prefer habitats 
situated at mid to high elevations (200-1341 m), up to 
the top of the islands in two cases. 


Distribution: Galapagos islands of Fernandina, Isabela, 
Pinta, Santa Cruz and Santiago; presumably endemic. 


Remarks: The vesica of two dissected males had no 
cornuti while another had seemingly lost only the 5 
shorter and thinner cornuti. 


ACKNOWLEDGEMENTS 


I thank the authorities of the Galapagos National Park 
for the collecting and export permits, the authorities 
and staff of the Charles Darwin Research Station, 
particularly Lazaro Roque-Albelo in 2002, 2004 and 
2005, for logistical support, and Stewart B. Peck 
(Carleton University, Ottawa) for inviting me to take part 
in his inventory of the Galapagos insects, funded by an 
operating grant of the Natural Sciences and Engineering 
Research Council of Canada to S. B. Peck for field 
research on arthropod evolution, for the organization of 
our 1989 and 1992 expeditions, and for being a model 
Ph.D. advisor and scientist. Furthermore, I wish to thank 
N. Castillo, C. Causton, J. Cook, M. Inca, J. Loaiza, 
R. Palma, L. Roque-Albelo, P. Schmitz, B. Sinclair, 
and E. Vilema for companionship and help in the field, 
the Galapagos Conservation Trust (London, U.K.) for 
financing my visit to the BMNH in 2000, the City of 


The Pyraustinae (Lepidoptera, Pyralidae s. 1.) of the Galapagos Islands, Ecuador 69 


Geneva for funding my 2002, 2004 and 2005 field trips, 
the Claraz, Lombard, and Schmidheiny Foundations as 
well as the Baslerstiftung ftir biologische Forschung, 
the Swiss Academy of Sciences, the City of Geneva, 
and the University of Geneva for grants awarded to 
P. Schmitz in 2004, 2005, and 2006, Martin Honey 
(BMNH) for dissecting and forwarding images of the 
slide preparation of a type specimen, Alan Tye and Frank 
Bungartz (CDRS) for information on plant distribution, 
J.-F. Landry for information on DNA bar-code data. For 
the loan of material in their care I am very grateful to 
D. Grimaldi, F. H. Rindge, and S. Rab-Green (AMNH), 
M. Shaffer and K. Tuck (BMNH), P. Arnaud, K. Ribardo 
and N. Penny (CAS), H. Herrera and L. Roque-Albelo 
(CDRS), J.-F. Landry (CNC), S. Cover, P. D. Perkins, and 
C. Vogt (MCZ), and the late John S. Garth (University of 
Southern California, Los Angeles, USA). Finally, I thank 
Philippe Wagneur (MHNG) for taking the photographs 
of the moths and for upgrading these images and others, 
Florence Marteau (MHNG) for skilfully producing the 
plates, and the reviewers of the manuscript for their 
constructive comments. 


REFERENCES 


Capps H.W. 1967. Review of some species of Loxostege Hübner 
and descriptions of new species (Lepidoptera, Pyraustidae: 
Pyraustinae). Proceedings of the United States National 
Museum, Washington 120 (3561): 1-75, 7 pls. 

Causton C.E., Peck S.B., Sinclair B.J., Roque-Albelo L., 
Hodgson C.J., Landry B. 2006. Alien Insects: Threats and 
Implications for Conservation of Galapagos Islands. Annals 
of the Entomological Society of America 99: 121-143. 

Guillermet C. 2009. Les Hétérocères, ou papillons de nuit, de 
l’île de La Réunion. Familles des Pyralidae et Crambidae. 
Association Nature, Découverte et Partage/ Parc National 
de La Réunion. 552 pp., 11 pls. 

Heppner J.B. 2003. Lepidoptera of Florida. Part 1. Introduction 
and catalog. Arthropods of Florida and neighbouring land 
areas 17: 670 pp. 

Jaramillo Diaz P., Guézou A. 2013. CDF Checklist of Galapagos 
Vascular Plants - FCD Lista de especies de Plantas 
Vasculares de Galapagos. In: Bungartz F., Herrera H., 
Jaramillo P., Tirado N., Jiménez-Uzcategui G., Ruiz D., 
Guézou A., Ziemmeck F. (eds). Charles Darwin Foundation 
Galapagos Species Checklist - Lista de Especies de 
Galapagos de la Fundacion Charles Darwin. Charles 
Darwin Foundation / Fundacion Charles Darwin, Puerto 
Ayora, Galapagos: http://\www.darwinfoundation.org/ 
datazone/checklists/vascular-plants/ Last updated 03 Jun 
2013. 

Kristensen N.P. 2003. Skeleton and muscles: adults (pp. 
39-131). Jn: Kristensen N. P., Lepidoptera, moths and but- 
terflies. Vol. 2: Morphology, physiology, and development. 
In: M. Fischer, Handbook of Zoology IV (36). Walter de 
Gruyter, Berlin and New York. 

Landry B. 2006. The Gracillariidae (Lepidoptera, Gra- 
cillarioidea) of the Galapagos Islands, Ecuador, with notes 
on some of their relatives. Revue suisse de Zoologie 113: 
437-485. 


Linsley E.G., Usinger R.L. 1966. Insects of the Galapagos 
Islands. Proceedings ofthe California Academy of Sciences, 
4th series, 33(7): 113-196. 

McMullen C.K. 1999. Flowering plants of the Galapagos. 
Cornell University Press, Ithaca and London, XIV + 
370 pp. 

Munroe E.G. 1976. Pyraloidea Pyralidae comprising the 
subfamily Pyraustinae tribe Pyraustini (part 1) [pp. 1-78, pls 
1-4, A-H; (conclusion); pp. 79-150, pls 5-9, J-U, pp. xiii- 
xvii]. /n: Dominick R. B. et al, The Moths of America 
north of Mexico 13.2A & 13.2B. E.W. Classey Ltd. and The 
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Munroe E.G. 1983. Pyralidae (except Crambinae) (pp. 67-76, 
78-85). In: Hodges R. W. et al., Check List of the Lepido- 
ptera of America north of Mexico including Greenland. £. 
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Foundation, London. 

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insect fauna of an oceanic archipelago: The Galapagos 
Islands, Ecuador. American Entomologist 44: 218-237. 

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17: 255-508. 


Revue suisse de Zoologie (mars 2014) 122(1): 71-74 


ISSN 0035-418 


A new species of the genus Matileortheziola Kozar & Foldi 
(Hemiptera: Coccoidea: Ortheziidae) 


M. Bora Kaydan!?, Zsuzsanna Konczné Benedicty! & Eva Szita!* 


! Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of Sciences, Budapest, Hungary 
Cukurova University, Imamoglu Vocational School, Adana, Turkey 


correspondent author: szita.eva@agrar.mta.hu 


Abstract: This paper describes a new species of the genus Matileortheziola Kozär & Foldi (Hemiptera: Coccoidea: 
Ortheziidae) from the Ethiopian region (Rwanda, Kenya). The specimens were extracted from forest litter using Berlese 
funnels, from the collections of the Muséum d’histoire naturelle de Genève, Switzerland. An identification key to the 


currently known species of Matileortheziola is provided. 


Keywords: Matileortheziolini - ensign scale - taxonomy 


INTRODUCTION 


Ensign scale insects (Hemiptera: Coccoidea: Orthezii- 
dae) are believed to be either ancestral to all scale insects, 
or a primitive, isolated branch of the archaeococcoids 
(Vea & Grimaldi, 2012). In the family there are two main 
groups of host-plant specialization; one group is com- 
posed of species that occur sporadically on leaf litter, 
mosses and lichens; the other feeds on vascular plants, 
including grasses, herbaceous and woody plants (Koteja, 
1996; Kozar, 2004; Vea & Grimaldi, 2012). 

The species of the subfamily Ortheziolinae are found 
in the Palaearctic, Oriental and Ethiopian regions, with 
the highest number of species occurring in the Ethiopian 
region. The subfamily is divided into four tribes: Orthe- 
ziolamametini, Ortheziolini, Matileortheziolini and 
Ortheziolacoccini (Kozar, 2004). The tribe Matile- 
ortheziolini is a monotypic tribe including only one genus, 
Matileortheziola Kozär & Foldi, which is characterised 
by having a narrow, compact wax plate band around the 
body margin and wide, plate-like structures on the mid- 
dorsum (Kozär & Foldi, 2000). The tribe is found only 
in the Ethiopian region and to date, only one species, 
Matileortheziola angolensis Kozär & Foldi, has been 
recorded (Kozär, 2004). This paper describes as new 
a second Matileortheziola species from the Ethiopian 
region, based on specimens extracted by Berlese funnels 
from forest litter. 


Manuscript accepted 04.08.2014 
DOI: 10.528 1/zenodo.14575 


- Ethiopian region. 


MATERIAL AND METHODS 


The specimens described and recorded in this study were 
all collected using soil and litter sampling devices, and 
were extracted by Berlese funnel from samples collected 
in the Ethiopian region. 

Specimens were prepared for light microscopy using the 
slide-mounting method discussed by Kosztarab & Kozar 
(1988). The morphological terminology used follows 
Kozar (2004). 

The type material of the new species here described is 
deposited in the Muséum d’histoire naturelle de Genéve 
collection (MHNG, with locality code, without registra- 
tion number) and in the Plant Protection Institute CAR 
HAS collection (PPI; with registration number). Mea- 
surements and count data of each character are given 
separately for the holotype, the data of paratypes and all 
the material available are given as a range in brackets. 


TAXONOMY 
Matileortheziola Kozar & Foldi, 2000 


Type species: Matileortheziola angolaensis Kozar & 
Foldi, 2000: 251. 


Diagnosis of the unmounted material (after Kozar, 
2004; with additions in order to accommodate the new 
species): Body whitish, segmentation not clearly visible. 
Only the margin of the dorsum covered with wax pro- 
trusions (some of which remain on the slide-mounted 
specimens); wax protrusions in some case circular in 


72 M. B. Kaydan, Z. Konezné Benedicty & È. Szita 


Key to species of Matileortheziola, based on adult females 


l Wax protrusions circular in shape; setae on legs and antennae spine-like; multilocular disc pores in a single row 


OMAMETOMEULSEIONOVISACID ar de 


RTS rasi. M. angolaensis Kozar & Foldi 


— Wax protrusions not circular in shape; setae on legs and antennae lanceolate; multilocular disc pores absent from 


ANTETONEULEIONOVISACIDAN TERRE 


shape, and including separate rows of spines from the 
earlier larval stage (Kozar, 2004). This kind of form and 
setal arrangement is not known to occur in other mem- 
bers of the family Ortheziidae. 


Diagnosis of the slide-mounted material: Slide- 
mounted adult female with antenna 3 segmented; third 
antennal segment bearing a slender apical seta, flagel- 
late sensory setae and small spine-like subapical setae; 
second segment with I campaniform sensillum. Eye 
stalk elongate, fused with pseudobasal antennal seg- 
ment. Legs well developed; leg setae robust, spine- 
like or lanceolate; trochanter and femur fused, tibia 
and tarsus fused; tibia with 1 campaniform sensillum 
and at least | fleshy sensory seta; tarsus without digit- 
ules; claw digitules spine-like, claw without a denticle. 
Labium two segmented, with 6-8 spine-like or lanceo- 
late; with 2 long setae near apex of labium very close 
together, each situated in single setal socket. Anal ring 
situated in a fold of derm on dorsal surface, bearing 6 
setae. Sclerotized plate present on dorsum anterior to 
anal ring, wider than long. A dorsal, narrow, compact 
wax plate band present around body margin. Wide plate- 
like structures present on mid-dorsum on the anal ring. 
Quadrilocular pores scattered over surface, appearing 
like microtubular ducts. Thumb-like pores present in 
a cluster lateral to anal ring. Abdominal spiracles ven- 
tral on anterior segments (segments I, II or III), with at 
least one present on each side; when present, posterior 
abdominal spiracles located on dorsum near anal ring, 
each surrounded by a cluster with or without multilo- 
cular disc pores. 


Matileortheziola lanceolata Kaydan 
& Konczné Benedicty sp. n. 
Fig. 1 


Material examined: Holotype MHNG, without regis- 
tration number; adult female; Rwanda (Kayove, 2100 m 
a.s.l.); leg. P. Werner, 15.v.1973 (MHNG locality code: 
Rwa-73/2; PPI code: 9610). — Paratypes: MHNG, with- 
out registration number; 4 females on 3 slides; Kenya 
(Kakamega, 1500 m a.s.l.); leg. L. Deharveng; 4.11.1974 
(MHNG locality code: KEN 12; PPI code: 9604). — PPI 
9606; 2 females on 2 slides; Kenya (Aberdare Nat. Park, 
2300 m a.s.l.); leg. V. Mahnert, 25.xi.1974 (MHNG 
locality code: KEN 6). 


Diagnosis: Matileortheziola lanceolata sp. n. a close 
relative of Matileortheziola angolaensis Kozar and 


TIRI RISI concour M. lanceolata sp. n. 


Foldi, but differs from this species by having (M. ango- 
laensis in parenthesis) (i) wax protrusions not circular in 
shape (wax protrusions circular in shape); (11) setae on 
legs and antennae lanceolate (setae on legs and anten- 
nae spine-like); (111) multilocular disc pores absent from 
anterior edge of ovisac band (multilocular disc pores in 
a single row on anterior edge of ovisac band). 


Etymology: The name of the species is derived from its 
unique lanceolate setae. 


Description of the slide mounted adult female: 

Body: 1.133 (0.927-1.133) mm long, 0.793 (0.700-0.824) 
mm wide. Antenna 3 segmented; eye stalk elongate, 
fused with pseudobasal antennal segment. Antennal seg- 
ment lengths as follows: Ist 60 (48-76) um; 2nd 41 (34- 
41) um; 3rd 146 (130-146) um. Second antennal segment 
with one campaniform sensilum; 3rd segment almost 
cudgel shaped (Fig. 1). Apical seta of antenna 84 (58-90) 
um long, the 2 subapical setae 29 (18-29) and 22 (11-22) 
um long respectively. Third antennal segment covered 
with 23-25 lanceolate setae, each about 12 um long. 
Venter: Labium two-segmented, 94 (72-100) um long. 
Stylet loop longer than labium. Anterior legs: coxa 60 
(60-72) um long, trochanter and femur 204 (156-206) 
um, tibia and tarsus 223 (170-223) um, claw 34 (29-34) 
um, claw digitules lanceolate, each 11 (10-11) um long; 
middle legs: coxa 78 (67-84) um, trochanter and femur 
216 (149-216) um, tibia and tarsus 228 (170-228) um, 
claw 34 (29-34) um, claw digitules lanceolate, each 11 
(10-12) um long; posterior legs: coxa 84 (77-92) um, 
trochanter and femur 240 (192-240) um, tibia and tarsus 
270 (228-270) um, claw 36 (31-36) um, claw digitules 
lanceolate, each 12 (10-13) um long. Claws each without 
a denticle. Legs with rows of lanceolate setae each 12 um 
long, and each leg with one sensory pore on tibia. Tho- 
racic spiracular openings each associated with a small 
marginal group of protrusions and pores and surrounded 
by small wax plates. Each spiracle with a pore cluster 
present at anterior margin. Anterior spiracle diameter 
18 (16-21) um. Thorax with scattered thick setae and 
quadrilocular pores, each 3-4 um in diameter. Abdomen 
with two rows of wax plates within ovisac band. A single 
row of multilocular disc pores each 4-5 um in diameter 
and with 6-12 loculi in the wax plates; also a row of mul- 
tilocular disc pores present anterior to vulva, each pore 
6-12 um in diameter. 

Dorsum: Dorsal wax plate only present in a narrow band 
along the margin; spines in wax plate each 12-14 um 
long. Wax plates with two kinds of setae: a truncate type, 


A new Matileortheziola sp. 


Fig. 1. Matileortheziola lanceolata Kaydan & Konczné Benedicty sp. n., holotype, adult female. 


74 M. B. Kaydan, Z. Konczné Benedicty & È. Szita 


each ca 16 um long; and long, slightly lanceolate, hair- 
like setae on edge of wax plate, each 12-14 um long. 
Submedian area of dorsum with four lanceolate setae, 
hair-like setae in a longitudinal row on each side. Derm 
wrinkled, covered with scattered quadrilocular pores 
each 2 um. Eight large, plate-like structures on mid-dor- 
sum of head, thorax and first four abdominal segments. 
Some circular quadrate pores also present near marginal 
wax plate row. Sclerotized anal plate present anterior to 
anal ring, 257 (200-257) um wide and 46 (46-90) um 
long, without setae. Anal ring 32 (30-34) um wide, 36 
(31-36) um long, containing two rows of pores and 6 anal 
ring setae, each 34 (28-34) um long. A group of thumb- 
like pores, each 3-5 um wide, present on each side of anal 
ring. Multilocular disc pores each 4-5 um in diameter and 
with 6-12 loculi, present around anal ring only. One pair 
of posterior abdominal spiracles present, situated on each 
side of anal ring. 


Remark: The unmounted adult female was not seen. 
Host plant: Unknown. 


Distribution: Rwanda, Kenya. 


ACKNOWLEDGEMENTS 


We would like to thank the OTKA (Hungarian National 
Science Found) (Grant No. T 048801, T 075889) for 
funding this project. The first author thanks TUBITAK 
for their financial support enabling study of the family 


Ortheziidae in Hungary. The authors wish to thank 
to Dr. Peter Schwendinger, the curator of Arachnida 
Collection of Muséum d’histoire naturelle de Genève, 
for his kind help and for making it possible for us to 
study the collections. Special thanks go Dr. Takumasa 
Kondo (Entomology Laboratory, CORPOICA, Pal- 
mira, Colombia), Dr. Gillian W. Watson (Plant Pest 
Diagnostic Laboratory, California Department of Food 
& Agriculture, Sacramento, California, USA) and Dr. 
Yair Ben-Dov (Department of Entomology, Agricultural 
Research Organization, Bet Dagan, Israel for reviewing 
the manuscript. 


REFERENCES 


Kosztarab M. & Kozar F. 1988. Scale Insects of Central Europe. 
Akademiai Kiado, Budapest, 456 pp. 

Koteja J. 1996. Scale insects (Homoptera: Coccinea) a day after 
(pp. 65-88). In: Schaefer, C. W. (ed.). Studies on Hemipte- 
ran Phylogeny. Thomas Say Publications in Entomology, 
Proceedings, Entomological Society of America, Lanham, 
Maryland. 

Kozar F. 2004. Ortheziidae of the World. Plant Protection Ins- 
titute, Hungarian Academy of Sciences, Budapest, 525 pp. 

Kozar F. & Foldi I. 2000. Matileortheziola angolaensis n. gen., 
n. sp. (Hemiptera, Coccoidea, Ortheziidae). Revue 
Frangaise d’Entomologie (N.S.) 22: 251-254. 

Vea I. & Gimaldi D. A. 2012. Phylogeny of ensign scale insects 
(Hemiptera: Coccoidea: Ortheziidae) based on the morpho- 
logy of recent and fossil females. Systematic Entomology 
37: 758-783. 


Revue suisse de Zoologie (mars 2015) 122(1): 75-120 ISSN 0035-418 


On the Scaphidiinae (Coleoptera: Staphylinidae) of the Lesser Sunda Islands 


Ivan Löbl 


Muséum d'histoire naturelle, Case postale 6434, CH-1211 Genève 6, Switzerland 
E-mail: ivan.lobl@bluewin.ch 


Abstract: The scaphidiines of the Lesser Sunda Islands are reviewed. Among the 45 species found within examined 
collections, following are described as new: Baeocera badia sp. nov., B. baliensis sp. nov., B. barda sp. nov., B. basalis 
sp. nov., B. batukogensis sp. nov., B. beata sp. nov., B. bella sp. nov., B. bifurcata sp. nov., B. bifurcilla sp. nov., B. bona 
sp. nov., B. brevis sp. nov., B. breviuscula sp. nov., Scaphisoma ablutum sp. nov., S. activum sp. nov., S. acutatum sp. 
nov., S. acutum sp. nov., S. adjunctum sp. nov., S. adscitum sp. nov., S. aequum Sp. nov., S. aereum sp. nov., S. affabile sp. 
nov., S. affectum sp. nov., S. angulare sp. nov., S. animatum sp. nov., S. antennarum sp. nov., S. approximatum sp. nov., 
S. aspectums p. nov., Scaphobaeocera baliensis sp. nov., S. lombokensis sp. nov., Scaphoxium bilobum sp. nov., Xoti- 
dium bolmarums sp. nov. Scaphisoma gracilicorne Achard, 1920, S. sapitense Pic, 1915 and Scaphobaeocera kraepelini 
(Pic, 1933) are redescribed. Scaphisoma sapitense infasciatum Achard, 1920 and S. dansalanense Löbl, 1972 are placed 
in synonymy of S. /uteomaculatum Pic, 1915. Lectotypes are designated for Scaphisoma gracilicorne Achard, 1920, 
S. luteomaculatum Pic, 1915, S. sapitense Pic, 1915, S. infasciatum Achard, 1920, S. testaceomaculatum (Pic, 1915), 
S. subelongatum (Pic, 1915) and Scaphobaeocera kraepelini (Pic, 1933). Keys to species of Baeocera, Scaphisoma, and 


Scaphobaeocera known from the Lesser Sundas, and a key to the world species of Xotidium are provided. 


Keywords: Coleoptera - Staphylinidae - Scaphidiinae - taxonomy - Lesser Sunda Islands. 


INTRODUCTION 


To date, only five species of the rove beetle subfamily 
Scaphidiinae have been reported from the Lesser Sunda 
Islands, 1.e., Sapitia lombokiana Achard, 1920, Scaphi- 
soma lombokense Löbl, 1986, S. sapitense Pic, 1915 (with 
its “variety” infasciatum Achard, 1920), S. sesaotense 
Löbl, 1986, and S. coarctatum Löbl, 1976). While Sapi- 
tia lombokiana is widely distributed in Southeast Asia, 
the Scaphisoma species are currently known only from 
Lombok (Löbl, 1997), with the exception of S. coarc- 
tatum which was recently reported from the Moluccas 
(Löbl, 2014). As some of the other islands of the archi- 
pelago are readily accessible and have, unless deforested, 
habitats convenient for sustainability of mycophagous 
beetles, a major gap was obvious. I have tried to fill it at 
least partly during a trip to Bali and Lombok, from the 
end of October to mid November 1991. My work was 
hampered by heavy rainfalls at the beginning of the mon- 
soon period, but other collections provided substantial 
complementary material. The study of the available col- 
lections yield 45 species in six genera that are reported 
below from Bali, Lombok, Sumbawa and Timor, among 
them 31 species are described as new. The absence of 
members of several genera, such as Cyparium Erichson, 


Manuscript accepted 27.08.2014 
DOI: 10.5281/zenodo.14582 


1845, Scaphidium Olivier, 1790, Bironium Csiki, 1909, 
and Scaphicoma Motschulsky, 1863, suggests still per- 
taining gaps in the knowledge of the group as far these 
islands concerned. The inadequate level of information is 
also suggested by the fact that many species are known 
only from a single locality or island. 


MATERIAL AND METHODS 


The material examined is housed in the following insti- 
tutes: Muséum d’histoire naturelle, Genève, Switzerland 
(MHNG), Muséum national d’Histoire naturelle, Paris, 
France (MNHN), Naturalis Biodiversity Center, Leiden, 
the Netherlands (NBCL), Narodni muzeum, Prague, 
Czech Republic (NMPC), and Staatliches Museum ftir 
Naturkunde, Stuttgart, Germany (SMNS). 

The locality data are given verbatim as on the respective 
labels, different labels are separated by a slash. The body 
length is measured from the anterior pronotal margin to 
the inner apical angle of the elytra. The maximal length 
and width ratios of the antennomeres are given, measured 
on slides. Characters given for metanepisterna concern 
their exposed parts. The abdominal sternites are counted 
from the first visible one (i.e., the third morphological). 


76 


The sides of the aedeagi refer to their morphological side, 
with the ostium situated dorsally, while it is in the resting 
position rotated 90°. Female characters are described and 
illustrated only in taxa for which they are discriminating. 
The eventually extruded parts of the internal sacs of the 
aedeagi are not considered in length measurements. Keys 
are based on well visible external characters, as far as 
possible. 


TAXONOMY 
Baeocera Erichson, 1845 


Comments: Baeocera is with over 260 species widely 
distributed and known from all continents but is poorly 


represented in northern temperate zones and absent 
from arid areas. Though Baeocera are often common 
in tropical and subtropical Asian forests, they are usu- 
ally under-represented in collections. Adequate sam- 
pling methods, such as sifting moist forest floor litter 
and using berlese or winkler devices for extraction of 
sampled insects, provide significant amount of spe- 
cimens. To date, only four species of Baeocera have 
been reported from the Indonesian Great Sunda Islands 
(Löbl, 1997) and three species from the Moluccas (Löbl, 
2014). The species from the Lesser Sundas are members 
of only two monophyletic group, the B. lenta group, 
which is in Asia particularly species-rich, and the B. ser- 
endibensis group. 


Key to the Baeocera species of the Lesser Sunda Islands 


Elytral punctation very fine and hardly visible at 100x magnification, similar to pronotal punctation, except on 
distinetlyapuneturediminuteslateromediantare AS PP ane B. bella sp. nov. 
= Elytral punctation distinct and much coarser than pronotal punctation on basal halves of disc at least, often on 
prevailing or entireidiscal! SUTFACE EE. rer ner eee TTT 2 
2 Metanepisterna distinct, usually wide, separated from metaventrite by distinct suture. Elytra with basal striae 
complete; joined to lateral Str ae 25.222285 ER eet 5 
— Metanepisterna concealed or very narrow, without distinct suture, lateral margin of metaventrite indicated by 
outer, eventually impressed, puncture row. Elytra usually with basal striae short, not joined to lateral striae, or basal 
Stride ADSENL i. AAA ROIO OTO. Ò 
3 Median part of sternite 1 punctate. Antennomere XI more than twice as long as wide. Parameres hardly curved at 
basetandino@wuldenedfapieallyzinWlateralkyi vv Ze B. badia sp. nov. 
— Median part of sternite 1 impunctate. Parameres distinctly curved at base and widened apically in lateral view...4 
4 Antennomere XI less than twice as long as wide. Internal sac of aedeagus with narrow, elongate, curved sclerite 
andsfinelyadenticulatehmem branes ER EEE B. barda sp. nov. 
= Antennomere XI more than twice as long as wide. Internal sac of aedeagus with robust, not curved sclerite and 
finelyzstriateimem branousistuucture SE B. baliensis sp. nov. 
D Elytra with sutural striae not curved at base, basal striae absent... B. bona sp. nov. 
= Elytra with sutural striae curved at base and extending laterally to form basal striae ................... nennen 6 
6 Basal halves, or prevailing surface of basal halves of elytra coarsely punctured, apical fourth to half of elytra 
becomingtabruptlyisMO0tN{OrgINCONSPICUOUSIVAPUNEIUTE (PRI EA TE etree 7 
_ Elytral punctation distinct on entire discal surface, or on almost entire disc, including areas near apical margins, 
eventuallyabecominoyoraduallyahineratow and A PIC ESP RER 9 
7 Elytraswithibasallsiniaezentires]o1neditopl a era [IST AE PR B. brevis sp. nov. 
_ Elytra with basal striae shortened, not reaching sides and not joined to lateral striae ....................... 8 
8 Antennomere VIII about 3 times as long as wide. Aedeagus with parameres widened in apical third and notched at 
levelfofitiprofemediantl obel(Gorsalivicw) Eee IT B. beata sp. nov. 
= Antennomere VIII slightly more than twice as long as wide. Aedeagus with parameres narrowed in apical third... 
naga aes RE RENI RIE DO ove UNS 20 TO OSATO ESE OI à Co CNTs I LOI III B. bifurcilla sp. nov. 
9 Elytratwithibasalistriae{completeSjoIneditollatera lista eee B. batukogensis sp. nov. 
= Elytra with basal striae shortened, not reaching sides and not joined to lateral striae ..................... 10 
10 Parameres slightly notched in middle part (dorsal view) and bent dorsally in apical part (lateral view). Tip of 
median lobe in level with parameral mid-length or reaching slightly beyond parameral mid-length........................ 
PIERRE E SR SE SE RER B. bifurcata sp. nov. 
= Barameresnotmotchedandnonbenttdorsall Mn api Al pe nn 11 
11 Apical process of median lobe short, with tip almost reaching apical third of parameral length. Internal sac without 
denticulate vesicle at level of apical part of sclerotized complex ....................... ri B. basalis sp. nov. 


Apical process of median lobe long, with tip reaching apical fourth of parameral length. Internal sac with denticulate 
vesiclelatilevellotapicalipartonsclerotizedieOMpIEx ee B. breviuscula sp. nov. 


Scaphidiinae of the Lesser Sunda Islands UO, 


Baeocera badia sp. nov. 
Figs 1-5 


Holotype: MHNG; £; Indonesia, Timor Camplong, 
250 m monsoon forest NE of village, leaf litt. D. Agosti, 
30.3.1991. 


Paratypes: MHNG; 4 &, 2 ©, 8 ex.; with the same 
data as the holotype (MHNG). — MHNG; 2 d, 5 ©, 
9 ex.; Timor NTT, Camping 25.3.91, leaf lit. D. Agosti 
F91598 [the latter site is NE von Cabang Luar Kota, 
10°1°57.57”S 123°56°4.04”E]. 


Etymology: The species epithet is a Latin adjective 
meaning chestnut-brown. 


Description: Length 1.45-1.50 mm, width 1.02- 
1.20 mm. Body moderately convex, without obvious 
microsculpture. Head and body reddish-brown, appen- 
dages and apex of abdomen lighter than body, almost 
yellowish. Length/width ratio of antennomeres as fol- 
lows: III 20/7: IV 24/7: V 30/7: VI 27/8: VII 34/9: VIII 
28/8: IX 34/15: X 32/15: XI 43/17. Lateral contours of 
pronotum and elytra separately arcuate. Pronotum with 
punctation very fine, setation hardly visible (50x magni- 
fication), lateral margins convex, anterior margin broad, 
basal lobe well developed; lateral pronotal carinae con- 
cealed in dorsal view. Tip of scutellum exposed. Elytra 
weakly narrowed apically, not covering abdominal 
apex, lateral margin carinae concealed or hardly visi- 
ble in dorsal view, sutural striae curved at base to form 
basal striae joined to lateral striae; adsutural areas flat 
near base, raised posteriad, parallel, punctate. Elytral 
impunctate or very finely punctate near base, with 
fairly coarse punctures on remaining surface, includ- 
ing apical areas, puncture intervals mostly as large to 
twice as large as puncture diameters. Epipleural striae 
reaching level of sternite 2, punctate, supraepipleural 
areas with puncture row. Hind wings well developed. 
Hypomera impunctate, smooth. Mesoventral suture 
distinct, mesoventrite distinctly punctate, with median 
ridge. Mesepimera large, each about 3 times as long as 
wide and about 3 times as long as interval to metacoxae. 
Median part of metaventrite flattened, with smooth cen- 
tre delimited laterally and apically by coarse punctures 
row. Lateral parts of metaventrite coarsely punctate, 
except on smooth areas near metacoxae. Punctures on 
lateral parts of metaventrite to part elongate, some as 
large as or larger than puncture intervals. Submesocoxal 
lines arcuate, with marginal punctures not extended 
laterally, submesocoxal areas about 0.05 mm long, as 
third of interval to metacoxae. Metanepisterna about 
0.06-0.07 mm wide, narrowed anteriad, with suture 
deep, broad, straight, coarsely punctate. Protibiae 
straight, mesotibiae and metatibiae slightly curved. 
Sternite 1 with coarse basal punctures uninterrupted in 
middle, separated by wrinkles about 0.02-0.08 mm long, 
remaining abdominal punctation distinct. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 


1-3, 5) 0.41-0.48 mm, fairly sclerotized. Median lobe 
symmetrical, with apical process shorter than basal bulb, 
weakly inflexed, gradually narrowed apically, with acute 
tip. Articular process distinct, not prominent. Parameres 
long, slightly narrowed from base to mid-length and 
somewhat bent in dorsal view, evenly narrow and straight 
in lateral view. Internal sac with curved, proximally 
widened sclerite and densely denticulate basal membra- 
nous lobe, membranes posterior sclerite extremely finely 
denticulate. 

Female: Ovipositor (Fig. 4) with long distal gonocoxite 
hardly narrowed apically, bearing two long apical setae, 
gonostyle present, with two short subapical setae and one 
long apical seta. 


Distribution: Indonesia: Timor. 


Type locality: Timor, Camplong, 250 m, monsoon 
forest NE of village. 


Comments: This is a member of the B. serendibensis 
group and is in external characters similar to B. ser- 
endibensis (Löbl, 1971) and B. sauteri Löbl, 1980. The 
aedeagal features suggest close relationships with the 
Australian B. alternans (Löbl, 1977), the New Guinean 
B. bironis (Pic, 1956), B. insperata (Löbl, 1977) and 
B. prospecta Löbl, 2002, and B. agostii Löbl, 2014 from 
the Moluccas. The new species may be distinguished 
from these species, B. agostii excepted, by the elytra 
having complete basal striae, joined to lateral striae, and 
all but B. prospecta by the distinctly punctate lateral 
parts of the metasternum. The shape of the sclerite of 
the internal sac and the absence of long, striate or hair- 
like structures in the proximal part of the internal sac are 
diagnostic for B. badia. 


Baeocera baliensis sp. nov. 
Figs 6-8 


Holotype: MHNG;  ; Indonesia, Bali Lake Tamblingan 
ca 1300 m, 30.X.91 I. Löbl, forest floor litt. 


Etymology: The species epithet is derived from the 
name of the island Bali. 


Description: Length 1.55 mm, width 0.96 mm. Body 
convex, lacking obvious microsculpture. Head, body 
and femora dark reddish-brown, tibiae slightly, tarsi 
and antennae distinctly lighter than body. Length/width 
ratio of antennomeres as follows: III 30/7: IV 30/7: V 
33/8: VI 25/6: VII 36/12: VIII 28/9: IX 35/14: X 32/15: 
XI 40/16. Lateral contours of pronotum and elytra con- 
tinuously arcuate. Pronotum with punctation very fine, 
setation hardly visible (50x magnification), lateral mar- 
gins convex, anterior margin broad, basal lobe poorly 
developed; lateral pronotal carinae concealed in dorsal 
view. Scutellum concealed. Elytra strongly narrowed 
apically, not covering abdominal apex, lateral margin 
carinae concealed in dorsal view, sutural striae curved at 


78 I. Löbl 


Figs 1-8. 
(1-5) Baeocera badia sp. nov., aedeagus in dorsal (1) and lateral (2) views, parameres (3) in ventral view, gonocoxite (4), internal sac 
in dorsal view (5). 


(6-8) Baeocera baliensis sp. nov., aedeagus in dorsal (6) and ventral (7) views, internal sac in dorsal view (8); scale bars for aedeagi 


= (0.1 mm, for parameres, gonocoxite and internal sac = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 79 


base to form basal striae joined to lateral striae; adsu- 
tural areas flat, parallel except in apical third, punctate. 
Elytra impunctate near base, with fairly coarse punc- 
tures on most surface, punctures becoming fine apically, 
puncture intervals mostly about as large to twice as 
large as puncture diameters. Epipleural striae reaching 
level of sternite 2, punctate, supraepipleural areas with 
puncture row. Hind wings well developed. Hypomera 
impressed, impunctate, smooth. Mesoventral suture 
distinct, mesoventrite impunctate, with median ridge. 
Mesepimera each about 3 times as long as wide and 
about 3 times as long as interval to metacoxae. Median 
part of metaventrite flattened, coarsely punctate except 
in middle part and near metacoxal process. Lateral parts 
of metaventrite coarsely punctate, except on smooth 
areas near metacoxae and metanapisterna. Punctures on 
lateral parts of metaventrite round, as large as or larger 
than puncture intervals. Submesocoxal lines arcuate, 
with marginal punctures not extended laterally, subme- 
socoxal areas about 0.06 mm long, slightly longer than 
half of interval to metacoxae. Metanepisterna about 
0.07 mm wide, parallel-sided, convex, with suture deep, 
broad, and coarsely punctate. Protibiae straight, meso- 
tibiae and metatibiae slightly curved. Sternite 1 with 
basal punctures fairly coarse, elongate, interrupted in 
middle, lacking basal wrinkles, punctation fine but dis- 
tinct on lateral parts of sternite, absent from median 
part. 

Male: Protarsomeres not widened. Aedeagus (Figs 6-8) 
0.35 mm long, fairly sclerotized. Median lobe similar to 
that in B. badia and B. barda, articular process almost 
indistinct. Parameres long, slightly narrowed behind 
mid-length and almost straight in dorsal view, bent in 
basal third and slightly widened in apically lateral view. 
Internal sac with robust, straight sclerite, membranes stri- 
ate laterally sclerite, denticulate structures absent. 


Distribution: Indonesia: Bali. 
Type locality: Bali, Lake Tamblingan, ca 1300 m. 


Comments: This species is likely closely related with 
B. praesignis Löbl, 2002 and B. bironis (Pic, 1956) 
that possess similar aedeagi, in particular the internal 
sacs. These two species may be, however, readily dis- 
tinguished from B. baliensis by the very finely punctate 
lateral parts of the metaventrite and by the compara- 
tively shorter apical antennomeres. 


Baeocera barda sp. nov. 
Figs 9-13 


Holotype: MHNG; d; Indonesia, Lombok Pusuk Pass 
300 m 3.X1.1991, I. Löbl degr. forest leaf litter. 


Paratypes: MHNG; 1 ©; with the same data as the 
holotype. - MHNG; 1 4; Lombok Bangko-Bangko 
(SW Lombok), 50 m degr. monsoonal for. D. Agosti 


19.3.1991. — SMNS; 1 €; Lombok Is. Senaro, N slope 
of Rinjani, 2-5.Feb 1994 Bolm Igt.1100 m. 


Etymology: The species epithet is a Latin adjective 
meaning slow. 


Description: Length 1.45-1.58 mm, width 0.87- 
1.05 mm. Colour and most external characters as in 
B. badia. Length/width ratio of antennomeres as fol- 
lows: II 17/7: IV 22/7: V 30/7: VI 26/8: VII 33/12: 
VIII 23/9: IX 30/13: X 28/15: XI 34/19. Mesepimera 
about 2.5 times as long as wide and 1.5 time as long 
as interval to mesocoxae. Punctation on lateral parts of 
metaventrite coarser than in B. badia. Submesocoxal 
lines strongly arcuate, submesocoxal areas about as long 
as half of interval to metacoxae. Metanepisterna not 
narrowed posteriad, suture straight, with variably large 
punctures. Sternite 1 impunctate in middle, basal punc- 
ture row interrupted between coxae. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
9-12) 0.37-0.43 mm long, fairly sclerotized. Median lobe 
similar to that in B. badia, articular process almost indis- 
tinct. Parameres long, slightly widened to mid-length 
and slightly sinuate in dorsal view, bent in basal third 
and slightly widened in apical third in lateral view. Inter- 
nal sac with almost evenly thick, curved and proximally 
tuberculate sclerite, basal membranous denticles well 
visible but distinct lobe absent. 

Female: Ovipositor (Fig. 13) with long distal gonocoxite 
slightly narrowed apically, bearing two long apical and 
four short subapical setae, gonostyle present, with one 
long apical seta. 


Distribution: Indonesia: Lombok. 
Type locality: Lombok, Pusuk Pass 300 m. 


Comments: This species is very similar to and obvi- 
ously closely related with B. badia. It may be reliably 
distinguished by its aedeagal characters, i.e., by the 
shape of the parameres and sclerotized piece of the 
internal sac. It differs from B. badia also by the impunc- 
tate centre of the sternite | and shorter apical antenno- 
meres. 


Baeocera basalis sp. nov. 
Figs 14-19 


Holotype: MHNG; ¢; Indonesia, Bali Mt. Agung above 
Besakih Temple, 1000-1100 m, 31.X.-1.X1.91, I. Löbl 
for.floor litter. 


Paratypes: MHNG; 3 2, 1 ©; with the same data as 
the holotype. - MHNG; 5 ©, 7 ©; Lombok Batu Koq 
(N of G. Rinjani) 500 m sec. forest in gorge D. Agosti 
12.03.1991. — MHNG; 3 4; Lombok Mt. Rinjani, ca 
400 m nr. Waterfalls, 5.XI.91 Löbl, veg.debris nr.river. 
—MHNG; 1 &, 1 ©; Lombok Gn. Rinjani 1000 m (win- 
kler) D. Agosti F91562. - MHNG; 1 <4, 2 ©; Lombok 


80 


I. Lobl 


5 


2 

{ 
IE 
tc 


NS 253% 
NN 


Figs 9-16 
(9-13) 


Baeocera barda sp. nov., aedeagus in dorsal (9) and lateral (12) views, parameres (10) in ventral view, tip of median lobe and 
paramere (11) in lateral view, internal sac (13) in dorsal view. 


(14-16) Baeocera basalis sp. nov., aedeagus in dorsal (14) and lateral (15) views, parameres (16) in ventral view; scale bars for 
aedeagi = 0.1 mm, for parameres, gonocoxite and internal sac = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 81 


Pusuk Pass, 300 m 3.X1.1991, I. Löbl degr. forest leaf 
litter. — 1 £; Lombok Mt. Rinjani above Senaro, 900- 
1100 m 6.XI.1991, I. Löbl forest floor litter. - MHNG; 
1 4; Lombok Batu Kog (N of G. Rinjani) 500 m sec. 
forest in gorge D. Agosti 12.03.1991. 


Etymology: The species epithet is a Latin adjective 
meaning basal. 


Description: Length 1.15-1.42 mm, width 0.79- 
0.95 mm. Body strongly convex, without obvi- 
ous microsculpture. Head and body very dark 
reddish-brown to blackish, appendages and apex of 
abdomen lighter than body. Length/width ratio of 
antennomeres as follows: III 25/7: IV 29/6: V 34/6: VI 
30/6: VII 37/8: VIII 30/7: IX 35/11: X 35/12: XI 40/15. 
Lateral contours of pronotum and elytra continuously 
arcuate. Pronotum with hardly visible punctation and 
setation (100x magnification), lateral margins convex, 
anterior margin broad, basal lobe well developed; lateral 
pronotal carinae concealed in dorsal view. Tip of scute- 
llum exposed. Elytra fairly strongly narrowed apically, 
not covering abdominal apex, lateral margin carinae 
concealed or hardly visible near base in dorsal view, 
sutural striae shallow, curved at bases to form basal 
striae abruptly ending about at mid-width of elytra; 
adsutural areas somewhat convex, parallel, very finely 
punctate. Discal punctation very fine near basal mar- 
gins and along lateral striae, fairly coarse about up to 
apical fourth, with punctures well delimited, puncture 
intervals about twice to four times as large as puncture 
diameters, becoming finer toward apices but still dis- 
tinct near apical margins. Epipleural striae entire, punc- 
tate, supraepipleural areas each with puncture row. Hind 
wings well developed. Hypomera impunctate, smooth. 
Mesoventral suture indistinct, mesoventrite punctate, 
lacking median ridge. Mesepimera about three times 
as long as wide and about twice as long as interval to 
metacoxae. Median part of metaventrite convex, smooth 
in middle, coarsely punctate around smooth centre. 
Entire lateral parts of metaventrite coarsely punctate, 
in particular near anterior margins, with punctures well 
delimited, mostly round, some punctures larges than 
puncture intervals. Submesocoxal lines arcuate, with 
marginal punctures not extending laterally, submeso- 
coxal areas about 0.03 mm long, about as third of inter- 
val to metacoxae. Metanepisterna fused to metaventrite, 
suture indicated by not impressed outer row of coarse 
punctures. Tibiae straight. Sternite 1 with basal puncture 
row uninterrupted in middle, lateral punctures elongate, 
not separated by wrinkles, finely but distinctly punctate 
posterior basal punctures. 

Male: Protarsomeres slightly widened. Aedeagus (Figs 
14-18) 0.30-0.34 mm long, moderately sclerotized. 
Median lobe symmetrical, with basal bulb large, longer 
than apical process, latter tapering, weakly inflexed, with 
acute tip and slightly concave ventral side (lateral view). 
Articular process moderately large. Parameres slightly 


bent posterior basal fourth and almost straight and evenly 
wide in lateral view, slightly narrowed and curved in api- 
cal halves in dorsal view. Internal sac with curved, almost 
evenly wide flagellum, flagellar guide-sclerite narrow, 
hook-like at base and blunt at apex, accessory sclerites 
absent, scale-like membranose structures extremely fine 
and hardly visible. 

Female: Ovipositor (Fig. 19) with distal gonocoxite long, 
narrowed apically, parallel-sided in apical section, bear- 
ing long apical seta, gonostyle absent. 


Distribution: Indonesia: Bali, Lombok. 


Type locality: Bali, Mt. Agung above Besakih Temple, 
1000-1100 m. 


Comments: This species is a member of the B. /enta 
group. Its aedeagal characters suggest relationship with 
B. louisi Löbl, 2012 from Luzon. The new species may 
be easily distinguished by the elytral punctation and 
sutural striae abruptly ending at elytral mid-width. 


Baeocera batukoqensis sp. nov. 
Figs 20-24 


Holotype: MHNG; <; Indonesia, Lombok Batu Koq 
(N of G. Rinjani) 500 m sec. forest in gorge D. Agosti 
12.03.1991. 


Paratypes: MHNG; 4 9; with the same data as the 
holotype. - MHNG; 1 £; Lombok Gn. Rinjani 1000 m 
(winkler) D. Agosti F91562. 


Etymology: The species epithet is derived from the 
name of the type locality. 


Description: Length 1.14-1.20 mm, width 0.67- 
0.72 mm. Body strongly convex, without obvious 
microsculpture. Head and body reddish-brown to black- 
ish-brown, hypomera, abdomen and appendages lighter 
than most of body. Length/width ratio of antennomeres 
as follows: III 27/6: IV 25/6: V 32/6: VI 33/6: VII 
40/8: VIII 32/8: IX 38/10: X 35/12: XI 35/15. Lateral 
contours of pronotum and elytra almost continuously 
arcuate. Pronotum with hardly visible punctation and 
setation (50x magnification), lateral margins convex, 
anterior margin broad, basal lobe small; lateral prono- 
tal carinae concealed in dorsal view. Tip of scutellum 
hardly visible. Elytra moderately narrowed apically, 
not covering abdominal apex, lateral margin carinae 
concealed or hardly visible near base in dorsal view, 
sutural striae shallow, not shortened, curved at bases 
to form basal striae extending laterally and joined to 
lateral striae; adsutural areas somewhat raised in mid- 
dle, parallel, impunctate. Discal punctation fairly 
coarse and dense from bases to mid-length or to apical 
third, with punctures well delimited, puncture intervals 
about as large to twice as large as puncture diameters; 
apical third to half of elytra finely but distinctly punc- 


I. Löbl 


SS assy 
nd SS, 


ES 
as 


3 


Figs 17-24 

(17-19) Baeocera basalis sp. nov., paramere (17) in lateral view, internal sac (18) in dorsal view, gonocoxite (19). 

(20-24) Baeocera batukogensis sp. nov., aedeagus in dorsal (20) and lateral (21) views, parameres (22) in lateral view, internal sac (23) 
in dorsal view, gonocoxite (24); scale bars for aedeagus = 0.1 mm, for parameres, internal sac and gonocoxite = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 83 


tate. Epipleural striae entire, punctate, supraepipleural 
areas each with puncture row. Hind wings well deve- 
loped. Hypomera impunctate, smooth. Mesoventral 
suture visible, mesoventrite finely punctate, lacking 
median ridge. Mesepimera about 2.5 times as long as 
wide and about twice as long as interval to metacoxae. 
Metaventrite coarsely punctate, punctures on lateral 
parts larger than on centre, to part larger than puncture 
intervals, only small central area and surfaces along 
metacoxae impunctate. Median part of metaventrite 
flattened. Submesocoxal lines arcuate, with marginal 
punctures not extending laterally, submesocoxal areas 
about 0.03 mm long, as third of interval to metacoxae. 
Metanepisterna fused to metaventrite, suture indicated 
by impressed outer row of coarse punctures. Tibiae 
straight. Sternite 1 with basal puncture row uninter- 
rupted in middle, basal punctures elongate and up to 
0.03 mm long laterally, punctation very fine, hardly visi- 
ble posterior basal puncture row. 

Male: Protarsomeres 1 to 3 widened. Aedeagus (Figs 
20-23) 0.42-0.44 mm long, fairly sclerotized. Median 
lobe symmetrical, with basal bulb large, longer than 
apical process, latter appearing short and wide in dorsal 
view, tapering, weakly curved, with tip bent and acute 
and ventral side oblique in lateral view. Articular pro- 
cess small. Parameres almost straight and posterior base 
evenly wide in dorsal view, slightly sinuate and evenly 
wide in lateral view. Internal sac with robust complex 
of sclerites, flagellum comparatively short, flagellar 
guide-sclerite narrow, small accessory rod joined to stri- 
ate membranous structures. 

Female: Ovipositor (Fig. 24) with long, gradually nar- 
rowed distal gonocoxite bearing long apical seta, gono- 
style absent. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Batu Koq N of G. Rinjani, 
500 m. 


Comments: This species is a member of the B. /enta 
group with aedeagal characters similar to those in 
B. papua (Löbl, 1975) from New Guinea, although the 
parameres are sinuate in the new species (lateral view). 
Baeocera batukogensis may be easily distinguished 
from B. papua by the abdominal ventrite 1 lacking dis- 
tinct basal wrinkles. 


Baeocera beata sp. nov. 
Figs 25-30 


Holotype: MHNG; &; Indonesia, Timor between Soe 
and Kapan, 1000 m evergreen for. on limest. D. Agosti, 
30.3.1991. 


Paratypes: MHNG; 3 ©; with the same data as the 
holotype. 


Etymology: The species epithet is a Latin adjective 
meaning fertile. 


Description: Length 1.15-1.17 mm, width 0.80- 
0.82 mm. Body strongly convex, without obvi- 
ous microsculpture. Head and body very dark 
reddish-brown to blackish-brown, appendages and apex 
of abdomen lighter than body. Length/width ratio of 
antennomeres as follows: III 25/7: IV 26/6: V 28/7: VI 
25/7: VII 30/7: VIII 25/8: IX 30/12: X 30/13: XI 35/15. 
Lateral contours of pronotum and elytra almost continu- 
ously arcuate. Pronotum with hardly visible punctation 
and setation (100x magnification), lateral margins con- 
vex, anterior margin broad, basal lobe well developed; 
lateral pronotal carinae concealed in dorsal view. Tip 
of scutellum hardly visible. Elytra weakly narrowed 
apically, not covering abdominal apex, lateral margin 
carinae concealed or hardly visible near base in dorsal 
view, sutural striae deep, not shortened, curved at bases 
to form basal striae abruptly ending about at mid-width 
of elytra; adsutural areas somewhat convex, parallel, 
impunctate. Discal punctation conspicuously coarse and 
fairly dense from bases to apical fourth, with punctures 
well delimited, puncture intervals about as large to twice 
as large as puncture diameters; apical fourth to third of 
elytra extremely finely punctate, appearing impunctate. 
Epipleural striae entire, punctate, supraepipleural areas 
each with puncture row. Hind wings well developed. 
Hypomera impunctate, smooth. Mesoventral suture 
indistinct, mesoventrite coarsely punctate, with median 
ridge hardly visible. Mesepimera about twice as long as 
wide and about twice as long as interval to metacoxae. 
Metaventrite all over coarsely punctate, punctures on 
lateral parts much larger than in middle, to part much 
larger than puncture intervals. Median part of metaven- 
trite flattened. Submesocoxal lines arcuate, with mar- 
ginal punctures not extending laterally, submesocoxal 
areas about 0.04-0.05 mm long, as half of interval to 
metacoxae. Metanepisterna fused to metaventrite, suture 
indicated by outer row of coarse punctures. Tibiae 
straight. Sternite 1 with basal puncture row uninter- 
rupted in middle, basal punctures elongate and separated 
by about 0.03-0.05 mm long wrinkles on lateral parts of 
sternite, posterior basal punctures appearing impunctate. 
Male: Protarsomeres hardly widened. Aedeagus (Figs 
25-28, 30) 0.34 mm long, moderately sclerotized. Median 
lobe symmetrical, with basal bulb large, longer than api- 
cal process, latter tapering, weakly curved, with acute tip 
and slightly concave ventral side (lateral view). Articular 
process moderately large. Parameres almost straight and 
distinctly widened in apical third, with shallow notch at 
level of tip of median lobe in dorsal view, bent in basal 
third in lateral view. Internal sac with gradually narrowed 
flagellum, lacking distinct guide-sclerite, with subbasal 
tubercle and membranes distinctly denticulate basally 
and apically. 

Female: Ovipositor (Fig. 29) with long distal gonoco- 
xite weakly narrowed apically, in apical section paral- 
lel-sided, bearing long apical seta, gonostyle absent. 


84 I. Löbl 


Figs 25-33 

(25-30) Baeocera beata sp. nov., aedeagus in dorsal (25) and lateral (26) views, parameres in dorsal (27) and lateral (28) views, 
gonocoxite (29), internal sac (30) in dorsal view. 

(31-33) Baeocera bella sp. nov., aedeagus in dorsal (31) and lateral (32) views, gonocoxite (33); scale bars for aedeagi = 0.1 mm, for 
parameres, internal sac and gonocoxite = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 85 


Distribution: Indonesia: Timor. 
Type locality: Timor, between Soe and Kapan, 1000 m. 


Comments: This species is a member of the B. /enta 
group. The aedeagal characters, in particular the shape 
of the parameres and the comparatively simple internal 
sac, suggest relationships with B. jankodadai Löbl, 2012 
from Palawan. The latter species differs by the compar- 
atively short antennomere VIII and the coarse elytral 
punctation not extending onto apical third of elytral 
disc. 


Baeocera bella sp. nov. 
Figs 31-33 


Holotype: MHNG; <; Indonesia, Timor Camplong, 
250 m monsoon forest NE of village, leaf litt. D. Agosti, 
BO SRIF 


Paratypes: MHNG; 3 ©, 8 ©; with the same data as the 
holotype. - MHNG; 5 €, 3 ©; Timor NTT, Camping 
25.3.91, leaf lit. D. Agosti F91598 [the site is NE von 
Cabang Luar Kota 10°1°57.57°S 123°56°4.04”E]. 


Etymology: The species epithet is a Latin adjective 
meaning pretty. 


Description: Length 0.92-1.08 mm, width 0.63- 
0.74 mm. Body strongly convex, without obvious 
microsculpture. Head and body light reddish-brown, 
appendages and apex of abdomen lighter than body, 
almost yellowish. Length/width ratio of antennomeres 
as follows: III 14/4.5: IV 14/4: V 18/4: VI 17/4: VII 
20/7: VIII 13/4.5: IX 17/10: X 17/12: XI 25/12. Lateral 
contours of pronotum and elytra continuously or almost 
continuously arcuate. Pronotum with hardly visible 
punctation and setation (100x magnification), lateral 
margins convex, anterior margin broad, basal lobe well 
developed; lateral pronotal carinae concealed in dorsal 
view. Tip of scutellum exposed. Elytra weakly narrowed 
apically, not covering abdominal apex, lateral margin 
carinae concealed or hardly visible near base in dor- 
sal view, sutural striae not shortened, curved at bases 
to form basal striae abruptly ending at mid-width of 
elytra; adsutural areas flat, parallel, impunctate. Elytral 
punctation very fine, as that on pronotum, except on 
fairly coarsely and densely punctate, about 0.10? to 
0.15? mm large lateromedian areas, and along lateral 
striae. Epipleural striae entire, impunctate, supraepi- 
pleural areas impunctate. Hind wings well developed. 
Hypomera impunctate, smooth. Mesoventral suture 
indistinct, mesoventrite very finely punctate, with 
median ridge hardly visible. Mesepimera large, each 
almost three times as long as wide and about 3 times as 
long as interval to metacoxae. Median part of metaven- 
trite somewhat flattened, with smooth centre delimited 
apically by distinct puncture row and laterally by fine 
punctation. Lateral parts of metaventrite distinctly 


punctate, usually smooth near metacoxae. Punctation 
on metaventrite consisting of round or slightly elon- 
gate punctures, mostly smaller than puncture inter- 
vals. Submesocoxal lines arcuate, with marginal 
punctures not extending laterally, submesocoxal areas 
about 0.03 mm long, as third of interval to metacoxae. 
Metanepisterna about 0.04-0.05 mm wide, paral- 
lel-sided, with deep, straight and punctate suture. Tibiae 
straight. Sternite 1 with basal wrinkles about 0.03-0.05 
mm long, abdominal punctation indistinct. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
31-32) 0.28-033 mm long, weakly sclerotized. Median 
lobe symmetrical, with basal bulb longer than apical 
process. Apical process inflexed, tapering, with acute 
tip and concave ventral side. Articular process indis- 
tinct. Parameres long, extending beyond tip of median 
lobe, weakly curved in dorsal and lateral views, each 
with fairly shallow notch at level of tip of median lobe. 
Internal sac with flagellum gradually narrowed, lacking 
flagellar guide-sclerite, with single wide and short acces- 
sory sclerite and membranous structure bearing minute 
denticles joined to its apex. 

Female: Ovipositor (Fig. 33) with long distal gonocoxite 
gradually narrowed apically, bearing single long apical 
seta, gonostyle absent. 


Distribution: Indonesia: Timor. 


Type locality: Timor, Camplong, 250 m, monsoon 
forest NE of village. 


Comments: This new species is a member of the 
B. lenta group and appears closely related with B. beata, 
though it differs drastically by the elytra with coarse 
punctation limited onto small, lateromedian area. The 
shape of the parameres in combination with the narrow 
flagellum is diagnostic for this new species. 


Baeocera bifurcate sp. nov. 
Figs 34-39 
Holotype: MHNG; ©; Indonesia, Bali Lake Buyan, ca 
1200 m 8.-9.X1.1991, I. Löbl degr. forest floor litter. 


Paratypes: MHNG; 1 ©; with the same data as the 
holotype. - MHNG; 1 4; Bali Badingkayu, 300-500 m 
10.-14.X1.1991, I. Löbl forest floor litter, bark. 


Etymology: The species epithet is a Latin adjective 
referring to the fork-like shape of the parameres. 


Description: Length 1.23-1.25 mm, width 0.88 mm. 
Body strongly convex, without obvious microsculpture. 
Head and body dark brown to blackish-brown, abdo- 
men, femora and tibiae lighter, tarsi and antennae yel- 
lowish. Length/width ratio of antennomeres as follows: 
IMI 23/5: IV 24/5: V 28/5: VI 25/5: VII 28/7: VIII 23/5: 
IX 33/10: X 34/12: XI 41/13. Lateral contours of prono- 
tum and elytra separately arcuate. Pronotum with hardly 


86 I. Löbl 


Figs 34-43 

(34-39) Baeocera bifurcata sp. nov., aedeagus in dorsal (34) and lateral (35) views, parameres in ventral (36) and lateral (37) views, 
internal sac (38) in dorsal view, gonocoxite (39). 

(40-43) Baeocera bifurcilla sp. nov., aedeagus in dorsal (40) and lateral (41) views, parameres in ventral (42) and lateral (43) views; 
scale bars for aedeagi = 0.1 mm, for parameres, internal sacs and gonocoxite = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 87 


visible punctation and setation (100x magnification), 
lateral margins convex, anterior margin broad, basal 
lobe small; lateral pronotal carinae concealed in dor- 
sal view. Minute tip of scutellum exposed. Elytra fairly 
narrowed apically, not covering abdominal apex, lateral 
margin carinae concealed or hardly visible near base in 
dorsal view, sutural striae curved at bases to form basal 
striae abruptly ending about at mid-width of elytra; 
adsutural areas flat, parallel, punctate. Elytral punctation 
very fine and similar to that on pronotum along basal 
margins and on apicolateral areas, coarse and densely 
punctate on remaining surface, many coarse punctures 
about as large as puncture intervals. Epipleural striae 
entire, punctate, supraepipleural areas impunctate. Hind 
wings well developed. Hypomera impunctate, smooth. 
Mesoventral suture indistinct, mesoventrite distinctly 
punctate, without median ridge. Mesepimera each about 
twice as long as wide and twice as long as interval to 
metacoxae. Median part of metaventrite somewhat 
flattened, with small smooth area in centre delimited 
by coarse punctation. Lateral parts of metaventrite 
coarsely punctate, except on narrow smooth areas near 
metacoxae; punctures not elongate, to part about as 
large as puncture intervals. Submesocoxal lines parallel, 
with marginal punctures not extending laterally, sub- 
mesocoxal areas about 0.02 mm long, about as fourth 
of interval to metacoxae. Metanepisternal suture indi- 
cated by outer row of coarse punctures. Tibiae straight. 
Sternite 1 with basal punctures uninterrupted in middle, 
coarse and slightly elongate on sides, basal wrinkles 
absent; punctation posterior basal puncture row distinct, 
becoming very fine apically. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
34-38) 0.34-0.35 mm long. Median lobe and parameres 
symmetrical, moderately sclerotized. Basal bulb of 
median lobe longer than apical process, latter inflexed, 
tapering, with acute tip and concave ventral side. Arti- 
cular process indistinct. Parameres conspicuously long, 
extending far beyond tip of median lobe, straight in dor- 
sal and lateral views, narrowed anterior level of tip of 
median lobe. Internal sac with complex basal sclerites 
forming arcuate flagellum and curved guide-sclerite 
joined to laterally expanded base. Membranes around 
apex of guide-sclerites forming small vesicle covered by 
scale-like structures. 

Female: Ovipositor (Fig. 39) with long distal gonocoxite 
strongly narrowed toward apical half, narrow and paral- 
lel-sided in long apical section, bearing single long apical 
seta, gonostyle absent. 


Distribution: Indonesia: Bali. 
Type locality: Bali, Lake Buyan, ca 1200 m. 


Comments: This species is a member of the B. lenta 
group and possibly allied with B. punctata (Löbl, 1975) 
from New Guinea. The shape of the sclerotized pieces 
of the internal sac are, however, diagnostic. In addition, 


most elytral punctures are in B. punctata larger than 
puncture intervals. 


Baeocera bifurcilla sp. nov. 
Figs 40-45 


Holotype: MHNG; <; Indonesia, Bali Yehbuah (N 
of Yehembang, E of Mendaya), 250 m D. Agosti 
25.4.1991. 


Paratypes: MHNG; 11 @, 12 ©; Indonesia, Bali 
Badingkayu, 300-500 m 10.-14.X1.1991, I. Löbl forest 
floor litter, bark. 


Etymology: The species epithet is a Latin noun, mean- 
ing small fork and referring to the shape of the para- 
meres. 


Description: Length 1.06-1.21 mm, width 0.72- 
0.80 mm. Body strongly convex, without obvious 
microsculpture. Head, body and femora reddish-brown, 
tibiae slightly lighter, tarsi and antennae yellowish. 
Length/width ratio of antennomeres as follows: III 
20/7: IV 16/7: V 22/7: VI 18/7: VII 30/8: VIII 16/7: 
IX 30/11: X 28/14: XI 32/14. Lateral contours of pro- 
notum and elytra separately arcuate. Pronotum with 
hardly visible punctation and setation (100x magnifi- 
cation), lateral margins convex, anterior margin broad, 
basal lobe small; lateral pronotal carinae concealed in 
dorsal view. Minute tip of scutellum exposed. Elytra 
weakly narrowed apically, not covering abdominal 
apex, lateral margin carinae hardly visible in dorsal 
view, sutural striae curved at bases to form basal striae 
abruptly ending about at mid-width of elytra; adsutural 
areas flat, parallel in anterior two thirds, impunctate. 
Elytral punctation very fine along sutural striae, evanes- 
cent along bases, on humeral areas and on apical third 
to two fifth. Punctation coarse and dense on remaining 
discal surface, with punctures well delimited, about 
half as large to as large as puncture intervals. Epipleural 
striae entire, punctate, supraepipleural areas impunc- 
tate. Hind wings well developed. Hypomera impunctate, 
smooth. Mesoventral suture indistinct, mesoventrite 
distinctly punctate, without median ridge. Mesepimera 
about 3 times as long as wide and 3 times as interval to 
metacoxae. Median part of metaventrite convex, with 
small smooth area in centre delimited laterally and pos- 
teriad by coarse punctation. Lateral parts of metaventrite 
coarsely punctate, except on narrow smooth areas along 
metacoxae; punctures round or slightly elongate, punc- 
ture intervals usually smaller than puncture diameters. 
Submesocoxal lines parallel, with marginal punctures 
not extending laterally, submesocoxal areas about 
0.02 mm long, about as fourth of interval to metacoxae. 
Metanepisternal suture indicated by impressed outer 
row of coarse punctures. Tibiae straight. Sternite 1 
with coarse basal punctures, separated by wrinkles up 
to about 0.05 mm long, uninterrupted in middle; punc- 


88 I. Löbl 


48 


Figs 44-52 

(44-45) Baeocera bifurcilla sp. nov., internal sac (44), gonocoxite (45). 

(46-50) Baeocera bona sp. nov., aedeagus in dorsal (46) and lateral (47) views, paramere with apical process of median lobe (48) in 
lateral view, internal sac (49) in dorsal view, gonocoxite (50). 

(51-52) Baeocera brevis sp. nov., aedeagus (51) and internal sac (52) in dorsal view; scale bars for aedeagi = 0.1 mm, for parameres, 
internal sacs and gonocoxite = 0.05 mm. 


Scaphidiinae of the Lesser Sunda Islands 89 


tation posterior basal puncture row distinct in middle, 
very fine laterally. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
40-44) 0.32-0.37 mm long. Median lobe and parameres 
symmetrical, moderately sclerotized. Basal bulb of 
median lobe longer than apical process, latter inflexed, 
tapering, with blunt tip and concave ventral side. Arti- 
cular process indistinct. Parameres long, extending fairly 
beyond tip of median lobe, straight in dorsal view, nar- 
rowed at level of tip of median lobe, arcuate in basal half 
and straight in apical half with slightly widened apical 
part in lateral view. Internal sac with complex basal 
sclerites forming arcuate flagellum and apically wide- 
ned guide-sclerite joined to laterally expanded base. 
Basal section of ejaculatory duct distinct. Membranes 
with extremely finely tubercle-like structures at base and 
around apex of guide-sclerites. 

Female: Ovipositor (Fig. 45) with long distal gonocoxite 
strongly narrowed toward apical half, narrow and paral- 
lel-sided in long apical section, bearing single long apical 
seta, gonostyle absent. 


Distribution: Indonesia: Bali. 


Type locality: Bali, Yehbuah (N of Yehembang, E of 
Mendaya), 250 m. 


Comments: This species is a member of the B. /enta 
group. Its aedeagal characters suggest close relation- 
ship with B. bifurcata. It may be easily distinguished by 
the shape of the parameres as seen in lateral view, the 
shorter apical section of the parameres, the shape of the 
flagellar guide-sclerite, and in external characters by the 
coarse elytral punctation abruptly ending before apical 
third of the disc. 


Baeocera bona sp. nov. 
Figs 46-50 


Holotype: MHNG; <; Indonesia, Lombok Batu Koq 
(N of G. Rinjani) 500 m sec. forest in gorge D. Agosti 
12.03.1991. 


Paratypes: MHNG; 3 &, 5 ©; Indonesia, Lombok 
Mt. Rinjani, ca 400 m nr. Waterfalls 5.XI.91 Löbl leg. 
debris nr. river. - MHNG; 2 ex.; Lombok, Mt. Rinjani 
above Senaro, 900-1100 m 6.XI.1991, I. Löbl forest 
floor litter. - MHNG; 1 9; Lombok Pusuk Pass 300 m 
3.X1.1991 I. Löbl degr. forest leaf litter. - MHNG; 3 3, 
1 9,9 ex.; Lombok Batu Kog (N of G. Rinjani) 500 m 
sec. forest in gorge D. Agosti 12.03.1991. — MHNG; 
5 ex.; Lombok, Gn. Rinjani 1000 m (Winkler) D. Agosti 
F91562. — SMNS; 1 ex.; Lombok Is.Senaro N slope of 
Rinjani, 2.-5.Feb. 1994 Bolm Igt. 1100 m. 


Etymology: The species epithet is a Latin adjective 
meaning good. 


Description: Length 0.98-1.13 mm, width 0.65- 


0.73 mm. Body rather strongly convex, lacking obvious 
microsculpture. Head and body reddish-brown, append- 
ages and apex of abdomen somewhat lighter than body. 
Length/width ratio of antennomeres as follows: III 22/5: 
IV 21/5: V 29/5: VI 27/6: VII 32/7: VIII 24/7: IX 33/10: 
X 31/11: XI 35/13. Lateral contours of pronotum and 
elytra separately arcuate. Pronotum with very fine punc- 
tation and setation usually distinct (50x magnification), 
lateral margins convex, anterior margin broad, basal 
lobe small; lateral pronotal carinae concealed in dorsal 
view. Scutellum concealed. Elytra weakly narrowed api- 
cally, usually covering abdominal apex, lateral margin 
carinae concealed or hardly visible near base in dorsal 
view, sutural striae shortened, starting posterior level 
on pronotal lobe; adsutural areas flat, parallel, punctate. 
Elytra with impunctate narrow lateroapical areas; ely- 
tral punctation coarse and dense, including along basal 
margins, puncture diameters on basal half about half as 
large to as large puncture intervals, becoming smaller 
apically. Epipleural striae entire, punctate, supraepi- 
pleural areas impunctate. Hind wings well developed. 
Hypomera impunctate, smooth. Mesoventral suture 
indistinct, mesoventrite distinctly punctate, median 
ridge absent. Mesepimera fairly large, each almost 3 
times as long as wide and about 3 times as long as inter- 
val to metacoxae. Median part of metaventrite slightly 
convex, with almost smooth centre delimited by coarse 
punctures. Lateral parts of metaventrite coarsely punc- 
tate, including near metacoxae. Punctation on metaven- 
trite well delimited, consisting of punctures round, 
mostly larger than puncture intervals. Submesocoxal 
lines arcuate, with marginal punctures not extending 
laterally, submesocoxal areas about 0.02 mm long, as 
fourth of interval to metacoxae. Metanepisternal suture 
indicated by outer puncture row. Tibiae straight. Sternite 
1 without basal wrinkles, basal punctures often elon- 
gate, forming row uninterrupted in middle, punctation 
near basal puncture row coarse, becoming very fine api- 
cally. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
46-49) 0.25-0.30 mm long. Median lobe symmetrical, 
with basal bulb longer than apical process. Apical process 
weakly inflexed, tapering, near tip very narrow and with 
concave ventral side (lateral view). Articular process 
small. Parameres long, extending beyond tip of median 
lobe, almost evenly broad, weakly curved in dorsal view, 
straight posterior basal third in lateral view. Internal 
sac with flagellum gradually narrowed, flagellar guide- 
sclerite weakly sclerotized, lacking accessory sclerite, 
membranous scale-like structures usually hardly visible. 
Female: Ovipositor (Fig. 50) with long distal gonocoxite 
strongly narrowed apically, in apical section parallel- 
sided, bearing long apical seta, gonostyle absent. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Batu Koq (N of G. Rinjani) 
500 m. 


90 


I. Löbl 


Figs 53-61 


(53-54) 
(55-59) 


(60-61) 


Baeocera brevis sp. nov., aedeagus (53) in lateral view, gonocoxite (54). 

Baeocera breviuscula sp. nov., aedeagus in dorsal (55) and lateral (56) views, internal sac (57) in dorsal view, paramere (58) 
in ventral view, gonocoxite (59); scale bars for aedeagi = 0.1 mm, parameres, internal sacs and gonocoxite = 0.05 mm. 
Scaphisoma ablutum sp. nov., aedeagus (60) in dorsal view; scale bar = 0.2 mm, (61) apical process of median lobe, with 
internal sac partly extruded; scale bar = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 91 


Comments: This species is a member of the B. /enta 
group. It may be readily distinguished from its 
Indonesian congeners, B. kaibesara Löbl from the 
Molucca Island Kai Besar excepted, by the shortened 
sutural striae of elytra. It shares with the latter species 
the shape of the median lobe and the parameres, differs 
however by the elytral punctation, coarser punctation on 
the ventrite 1, and by the shape of the sclerotized pieces 
of the internal sac. 


Baeocera brevis sp. nov. 
Figs 51-54 


Holotype: MHNG; &; Indonesia Bali Mt. Batukaru 
near Luhur Temple, 500-700 m, I. Löbl, 28.-29.X.1991. 


Paratypes: MHNG; 6 €, 6 9, 11 ex.; with the same 
data as the holotype. - MHNG; 1 4, 1 © Bali Batukaru 
18.VI.84 Rougemont. 


Etymology: The species epithet is Latin adjective 
meaning short and referring to the short distal gonoco- 
xite. 


Description: Length 0.98-1.13 mm, width 0.65- 
0.73 mm. Body rather strongly convex, lacking obvi- 
ous microsculpture. Head and body dark reddish-brown 
to blackish-brown, femora and tibiae somewhat lighter 
than body, tarsi and antennae distinctly lighter, yellow- 
ish. Length/width ratio of antennomeres as follows: III 
28/6: IV 30/6: V 34/5: VI 37/5: VII 45/7: VIII 37/6: IX 
45/9: X 42/12: XI 44/12. Lateral contours of pronotum 
and elytra separately arcuate. Pronotum with punctation 
and setation very fine, usually distinct (SOx magnifica- 
tion); lateral margins convex, anterior margin broad, 
basal lobe small; lateral pronotal carinae concealed in 
dorsal view. Scutellum concealed. Elytra fairly nar- 
rowed apically, not covering abdominal apex, lateral 
margin carinae concealed in dorsal view, sutural striae 
curved at base to form complete basal striae, joined to 
lateral striae; adsutural areas flat, parallel, punctate. 
Basal halves of elytra with punctation coarse and dense, 
coarse punctures usually also along basal and lateral 
margins; puncture diameters mostly about as large as 
puncture intervals. Apical halves of elytra about as 
finely punctate as pronotum. Epipleural striae entire, 
punctate in basal halves, supraepipleural areas impunc- 
tate. Hind wings well developed. Hypomera impunctate, 
smooth. Mesoventral suture indistinct, mesoventrite 
distinctly punctate, median ridge absent. Mesepimera 
fairly large, each almost 3 times as long as wide and 
about 3 times as long as interval to metacoxae. Median 
part of metaventrite slightly convex, almost entirely 
coarsely punctate. Entire lateral parts of metaventrite 
covered by coarse punctures larger than those on cen- 
tre of metaventrite. Punctures on metaventrite well 
delimited, round, mostly larger than puncture intervals. 
Submesocoxal lines arcuate, with marginal punctures 
hardly extending laterally, submesocoxal areas about 


0.02-0.03 mm long, as fifth to fourth of interval to 
metacoxae. Metanepisternal suture indicated by some- 
what impressed outer puncture row. Tibiae straight. 
Sternite 1 without basal wrinkles, basal punctures usu- 
ally elongate, uninterrupted in middle, and laterally usu- 
ally 0.02-0.04 mm long, punctures posterior basal row 
very fine, similar to those on apical parts of elytra. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
51-53) 0.39-0.44 mm long. Median lobe symmetrical, 
with basal bulb much longer than apical process. Api- 
cal process strongly inflexed, tapering, with ventral side 
oblique (lateral view). Articular process small. Parameres 
long and comparatively broad, extending far beyond 
tip of median lobe, almost evenly wide in basal halves, 
narrowed and weakly curved in level of tip of median 
lobe (dorsal view), almost straight and weakly widened 
toward level of tip of median lobe in lateral view. Internal 
sac with flagellum robust, sinuate, gradually narrowed, 
flagellar guide-sclerite strongly sclerotized, lacking 
accessory sclerite, membranous scale-like structures usu- 
ally hardly visible. 

Female: Ovipositor (Fig. 54) with distal gonocoxite 
short, strongly and gradually narrowed apically, bearing 
long apical seta, gonostyle absent. 


Distribution: Indonesia: Bali. 


Type locality: Bali Mt. Batukaru near Luhur Temple, 
500-700 m. 


Comments: This species is a member of the B. lenta 
group. The aedeagal characters, in particular the shape 
of the wide parameres and the short, strongly inflexed 
apical process of the median lobe, suggest relationships 
with B. carinata (Löbl, 1975) from Sumatra. The scle- 
rotized pieces of the internal sac are, however, distinc- 
tive in these two species. 


Baeocera breviuscula sp. nov. 
Figs 55-59 


Holotype: MHNG; €; Indonesia, Bali Badingkayu, 
300-500 m 10.-14.XI.1991, I. Löbl forest floor litter, 
bark. 


Paratypes: MHNG; 5 €, 7 ©; with the same data as 
the holotype. - MHNG; 1 £; Lombok Mt. Rinjani, ca 
400 m nr. Waterfalls, 5.XI.91 Löbl veg.debris nr.river. 


Etymology: The species epithet is Latin adjective 
meaning short and referring to the comparatively short 
parameres. 


Description: Length 0.95-1.07 mm, width 0.61- 
0.77 mm. Body strongly convex, without obvi- 
ous microsculpture. Head, body and femora light 
reddish-brown, tibiae slightly lighter, tarsi and antennae 
yellowish. Length/width ratio of antennomeres as fol- 
lows: III 18/6: IV 17/5: V 25/5: VI 23/5: VII 28/8: VII 


92 I. Löbl 


23/6: IX 30/10: X 30/13: XI 33/13. Lateral contours of 
pronotum and elytra separately arcuate. Pronotum with 
hardly visible punctation and setation (100x magnifi- 
cation), lateral margins convex, anterior margin broad, 
basal lobe small; lateral pronotal carinae concealed in 
dorsal view. Scutellum concealed. Elytra weakly nar- 
rowed apically, not covering abdominal apex, lateral 
margin carinae hardly visible in dorsal view, sutural 
striae curved at bases to form basal striae abruptly 
ending about at mid-width of elytra; adsutural areas 
flat, parallel in anterior two thirds, impunctate. Elytral 
punctation fairly coarse on most of anterior two thirds 
of disc, evanescent on small humeral area and becom- 
ing very shallow and fine from middle third toward 
apex; coarse punctures well delimited, mostly about 
half as large as puncture intervals. Epipleural striae 
entire, punctate, supraepipleural areas impunctate. 
Hind wings well developed. Hypomera impunctate, 
smooth. Mesoventral suture indistinct, mesoventrite 
distinctly punctate, without median ridge. Mesepimera 
large, about 3 times as long as wide and 3 times as long 
as interval to metacoxae. Median part of metaventrite 
flat, all over coarsely punctured. Entire lateral parts 
of metaventrite coarsely punctate; punctures round 
or hardly elongate, notably smaller and denser near 
metacoxae than near mesepimera, puncture intervals 
usually smaller than puncture diameters. Submesocoxal 
lines parallel, with marginal punctures not extend- 
ing laterally, submesocoxal areas about 0.02 mm long, 
about as fifth of interval to metacoxae. Metanepisternal 
suture indicated by impressed outer row of coarse punc- 
tures. Tibiae straight. Sternite 1 with basal punctures 
coarse, not or weakly elongate, up to about 0.02 mm 
long, not separated by wrinkles and uninterrupted in 
middle; punctation strongly reduced posterior basal row, 
usually consisting of few fine punctures. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
55-58) 0.28-0.32 mm long. Median lobe symmetrical, 
with basal bulb moderately longer than apical process. 
Apical process strongly process weakly inflexed, taper- 
ing, with most of ventral side almost oblique, tip bent 
and narrow (lateral view). Articular process fairly large. 
Parameres comparatively broad and short, moderately 
extending beyond tip of median lobe, almost evenly wide 
between basal third and apices in dorsal view, slightly 
widened apically in lateral view. Internal sac with fla- 
gellum narrow, arcuate, gradually narrowed, flagellar 
guide-sclerite weakly sclerotized, joined to basal com- 
plex of sclerites, membranous scale-like structures usu- 
ally distinct. 

Female: Ovipositor (Fig. 59) with long distal gonocoxite 
gradually narrowed apically, bearing long apical seta, 
gonostyle absent. 


Distribution: Indonesia: Bali, Lombok. 


Type locality: Bali, Badingkayu, 300-500 m. 


Comments: This species is a member of the B. lenta 
group. In external characters it resembles B. bifurcilla 
from which it may be distinguished by the elytra dis- 
tinctly punctured near pronotal lobe and on apical third, 
the median part of the metaventrite flat and entirely 
coarsely punctate, and the sternite 1 with basal punc- 
tures not or weakly elongate. 


Sapitia Achard, 1920 


The genus includes three South-East Asian species, all 
reported also from Indonesia though only one of them 
from the Lesser Sundas. Members of Sapitia are presum- 
ably associated with termites (Leschen & Löbl, 2005). 


Sapitia lombokiana Achard, 1920 


Distribution: Indonesia: Lombok, Sumatra; Vietnam; 
Philippines: Palawan (doubtful record). 


Comments: The species was not represented in collec- 
tions recently examined. Its description was based on 
specimens from Lombok: Sapit. Diagnostic characters 
including illustrations of the aedeagus, the synonymy 
with Baeoceridium sericeum Pic, 1922, and additional 
records are given in Löbl, 1978. 


Scaphisoma Leach, 1815 


Comments: Scaphisoma is with over 600 species cur- 
rently recognized as valid the most species-rich genus of 
the subfamily Scaphidiinae. Not surprisingly, it is with 
24 encountered species also quite diverse in the Lesser 
Sunda Islands. All have well developed hind wings sug- 
gesting dispersal ability and possible wide distribution. 


Scaphisoma ablutum sp. nov. 
Figs 60-62 


Holotype: MHNG; d; Indonesia, Indo: Lombok G. 
Rinjani 1950 m mossforest D. Agosti 16.3.1991 / Agosti 
F91558 Lombok G. Rinjani 8 16.3.91 [handwritten]. 


Paratypes: SMNS, MHNG; 2 ©; Indonesia, Lombok 
Is. Sapit-Sembalun Bumbung 14.-16.Feb.1994 Bolm 
Igt., 900-1500 m. — SMNS; 1 ©; Indonesia Lombok 
Serano N. Slope of Rinjani, 2.-5.Feb 1994 Bolm Igt. 
1100 m. 


Etymology: The species epithet is a Latin adjective, 
meaning washed and referring to a heavy rainfall in 
Lombok. 


Description: Length 1.90 mm, width 1.23 mm. Head 
and most of body light brown, apical third of elytra 
yellowish. Appendages somewhat darker than apical 
part of elytra. Pronotum and elytra not microsculp- 
tured. Antennae long, length/width ratio of antenno- 


Scaphidiinae of the Lesser Sunda Islands 93 


Key to the Scaphisoma species of the Lesser Sunda Islands 


I 


IRON RC] 


N 


10 


11 


12 


13 


14 


15 


16 


19 


20 


Elytra with sutural striae conspicuously strongly converging apically .…............................................................ 2 
Elytra with sutural striae parallel, or to large extent parallel, or weakly converging apically ................................. 4 
Pronotum and elytral disc with similar, very fine punctation. Pronotum and most of elytra very dark reddish-brown 
tolblacksapicaletounthrotachy easy e lO vs De S. jacobsoni Löbl 
Elytral disc with punctation much coarser than pronotal punctation. Colour pattern of pronotum and elytra 
CLT CTEM nn nn esse on ans en sec n ste Lucu sert ou rs ste sente tee eue aseeeorrenton tentes 3 
Pronotum bicoloured in male, dark brown to black in female. Elytra dark with light subhumeral spot and light 
subapical fascia. Antennomere III subcylindrical, elongate ........................... 220000 S. aspectum Sp. nov. 


Pronotum light, yellowish or reddish-brown in both sexes. Elytra light on prevailing surface, darkened along 
suture, basal margins and on transverse band posterior mid-length. Antennomere III subtriangular, short .............. 


een terso ars sic seras sons aneti sacesiarssuassecsssiioviesvnrsevnsseatevssoosseseseseseseseesess S. dohertyi Pic 
Elytra light on prevailing surfaces, darkened along lateral margin, base and sutural striae, each elytron with small, 
‚golmisil, cenz dal SPORE I OT S. sapitense Pic 
@olourgpatterno REINA TECHE e e II nn een anses ceceno 5 
Elytra with sutural striae extending along basal margins to form basal striae ................... 6 
Elytra with sutural striae not extending along basal margin and without basal striae...................... ii 8 
Antennomere V longer than III and IV combined, and much shorter than antennomere VIII ..S. acutatum sp. nov. 
Antennomere V as long as III and IV combined, almost as long as antennomere VIII ........................................ 7 
Body 1.55-1.60 mm long, uniformly reddish brown, elytra with shallow basal striae .............. S. lombokense Löbl 
Body 1.90 mm long. Pronotum dark reddish-brown, elytra coloured as pronotum along bases, apices, sutural striae 
and lateral margin, most of elytra much lighter, forming large reddish spot... Scaphisoma sp. 
Basomedian area of sternite 1 coarsely punctate. Metaventrite with antecoxal puncture rows. Elytra coarsely 
punctate, light in apical third, with sutural striae abruptly obliquely bent near base.................. S. affectum sp. nov. 
Rasomnedian part ot siano I verziimaly MINA PP en 9 
Metaventritehwithti ed ANIPIOONE SIRO O e ti NESSO Nieto cinici 10 
IMEtAVENtI TERI (NO UIMECIANILIOOVE SRO e ottieni entire iii 13 
EIVITATANAPTONOLUMIUNIFOTMIVACO OUTEA EEE O enni eh tes sne ee dns tres me es en 11 
Apical third of elytra clearly lighter than remaining elytral surface and pronotum ................. i 12 


Pronotum and elytra reddish-brown. Submesocoxal areas about as long as submetacoxal areas S. aequum sp. nov. 
Pronotum and elytra blackish-brown to black. Submesocoxal areas clearly longer than submetacoxal areas 


RE RA EEE OER entente tem ee sentent WaoneesaGinedsabisestocsdenesioabeeescvans S. aereum Sp. nov. 
AntennomereslfandiVAcombIMESNOTLEMNANN EE RI n S. adjunctum Sp. nov. 
AntennomeresgliltandiVAcomb Ned, on Sertha VAS RE O RIO S. ablutum sp. nov. 
Submetacoxal areas conspicuously large, extending onto apical half of sternite.................. S. antennarum sp. nov. 
Submetacoxalfareasnogextending{ontoapicalthalito NSTENNILE Een 14 
Hypomera with striate microsculpture. Anterior two thirds of elytra with conspicuous coarse punctation, apical 
thirditolwomitthvotielytraismoothwappeanin EM punctate ee. S. rufescens (Pic) 
Ely pomeranogmierosculprured SE ly traliPune ta ONCE] IE ERE O SPOSO Ono 13 
IMedian pari alnneaveninie wii SARDA TEOR MITE PR IAN 16 
IVIECAVENTE TEMO MICIOSCUIPIULCA RR RE nr nni 19 
Median part of metaventrite much coarser punctate than lateral parts, or than sternite 1, metaventrite with distinct 
INIECOXAPUNCHWICHOWSPRET ARE O Sentina S. sesaotense Löbl 
Median and lateral parts of metaventrite with similar, very fine punctation ................... 17 
Larger species 1.93-2.10 mm long. Elytra black, somewhat lighter along base and on adsutural areas and with 
Narrowlypy.clLOwISHYApI CES I RIE SIT ciro cene den onnesnme ec 40e S. angular Sp. nov. 
Smallemspecies MAS ECO MMIONLACOOMIPALETNO NEVA ILE Ron en 18 
Internal sac of aedeagus with mesal row of large, triangular sclerites, long lateral denticles fused to form plates and 
tuft of apical denticles covered by membranous spine-like structures....................... S. luteomaculatum Pic 
Internal sac of aedeagus with few long lateral sclerites, lacking mesal row of triangular sclerites............................ 
CA ITITTANIITIT ILARIA SCIITA DIVIETI S. obliquemaculatum Motschulsky 
BIKAR ad TETI WII Sima ere, ven7iime PISO oco000000000000000000000000000006008000060050500005000025000005 0050955058 IAA 20 
Elytral punctation distinctly coarser than pronotal punctation, eventually near bases similar to pronotal, on most 
SUTTACE ICO ATSC EP ee eecrentstsnesossesssnsenetesnessanestsstsssentnenhentnnestunennenten 21 
Antennomere IV about twice as long as III, combined with III about as long as V.................... S. coarctatum Löbl 


Antennomere IV about three times as long as III, combined with III distinctly longer than V ... S. affabile sp. nov. 


94 


N | 
(N°) 


N | 
Uo 


I. Löbl 


Pronotum and elytra on basal part of disc and on transverse subapical band dark brown or reddish-brown, most 
of elytral disc and narrow apical band light, yellowish. Submetacoxal areas large, almost reaching sternal mid- 
length: rs are OI et soe renee outs Sonar oe a eer ence S. animatum sp. nov. 
Blytralicolounditicrentasubmetacoxalfancas pm ano CT 22 


Sternite | about as metaventrite very finely punctate, submetacoxal lines COnvex................ 23 
Elytral punctation fine, in particular on basal halves, consisting of poorly delimited punctures. Apical two fifth 
somewhaglichterathtanibasalttWwo}fifthofCy{rA Re I S. gracilicorne Achard 
Elytral punctation distinct, fairly coarse, consisting of well delimited punctures. Elytra becoming lighter only near 
apical'margins...;-onartial ra PE TORE ATRIA I TI 24 
Antennomere IV about 4.5 times as long as III. Submetacoxal lines reaching sternal mid-length............................ 
car RR RR IT NE S. adscitum sp. nov. 
Antennomere IV about 2.5 times as long as III. Submetacoxal lines not reaching sternal mid-length.................. 25 
Aedeagus with wide apical process and bases of parameres strongly widened. Elytral punctation very fine along 
lateralimarginsSfairly{coarsetonimosHofidiscaliSUrta CCR S. activum Sp. nov. 
Aedeagus with narrow apical process and bases of parameres moderately widened. Elytral punctation fairly coarse, 
includingfonisurfacestalongalatera Marg SA O ORI S. acutum Sp. nov. 


meres as follows: III 16/9: IV 50/8: V 55/9: VI 55/9: 
VII 60/13: VII 50/10: IX 56/15: X 55/15: XI 65/17. 
Antennomere III subtriangular. Pronotum finely punc- 
tate, punctures dense, well delimited, lateral margins 
evenly rounded, lateral margin carinae not visible in 
dorsal view. Point of scutellum exposed. Elytra with 
lateral margin carinae not visible in dorsal view, apical 
margins truncate, inner apical angle rounded, not pro- 
minent, situated posterior level of outer angles, sutural 
margin raised, sutural striae fairly deep, converging api- 
cally, not curved at base and not extending laterad pro- 
notal lobe, adsutural areas flat. Elytral punctation fine 
and dense, punctures fairly well delimited, puncture 
intervals mostly about twice to three times as large as 
puncture diameters. Hypomera smooth. Mesepimeron 
about 1.5 times as interval to mesocoxa, about 4 times 
as long as wide. Metaventrite not microsculptured, flat 
in middle, with deep mesal groove and two shallow api- 
comedian impressions; punctation even, very fine and 
sparse. Submesocoxal areas 0.09 mm, almost as half of 
interval to metacoxa, submesocoxal lines convex, with 
fairly coarse marginal punctures. Metanepisternum flat, 
in level with metaventrite, not narrowed anteriad, with 
inner margin straight, rounded at angles. Tibiae straight. 
Abdomen very finely punctate, with striate microsculp- 
ture. Submetacoxal areas 0.04 mm, submetacoxal lines 
convex, with fine margin punctures. 

Male: Tarsomere | of protarsi and mesotarsi conspicu- 
ously widened, wider that tibiae, with conspicuous tenent 
setae. Tarsomeres 2 of protarsi and mesotarsi distinctly 
widened, tarsomeres 3 slightly widened. Aedeagus 
(Figs 60-62) 0.70 mm long, asymmetrical. Median lobe 
with basal bulb weakly sclerotized, robust and promi- 
nent articular process excepted. Apical process strongly 
inflexed, almost tubular, hardly narrowed apically in dor- 
sal view. Parameres fairly wide near bases, not lobed, 


bent and narrowed in apical halves in dorsal view, sinuate 
and slightly narrowed apically in lateral view. Internal 
sac with curved flagellum and membranes bearing spine- 
like structures. 


Distribution: Indonesia: Lombok. 
Type locality: Lombok, G. Rinjani 1950 m. 


Comments: The median lobe with robust articular pro- 
cess, the asymmetrical apical process, and the presence 
of a flagellum suggest relationships with members of 
the S. unicolor group, though the spine-like structures of 
the internal sac are absent in membersof that group. The 
short and robust median lobe in S. ablutum resembles 
that in the New Guinean S. fasciatum Pic, 1956 while 
the shape of the parameres is unique. See also com- 
ments under S. adjunctum. 


Scaphisoma adjunctum sp. nov. 
Fig. 63-65 


Holotype: SMNS; 4; Indonesia, Lombok Is. Senaro N 
slope of Rinjani 2.-5.Feb.1994 Bolm Igt. 1100 m. 


Paratypes: SMNS, MHNG; 2 9; with the same data as 
the holotype. - MHNG; 2 ©; Lombok Mt. Rinjani ca 
400 m nr. Waterfalls, 5.XI.91 Löbl, veg. debris nr. river. 
— MHNG; 2 ©; Indo: Lombok Batu Koq-G.Rinjani 
1200 m, sec. montane for. on slope, in ravine D. Agosti, 
133.01. 


Etymology: The species epithet is a Latin adjective, 
meaning added. 


Description: Length 1.42-1.60 mm, width 0.92-1.02 
mm. Head and most of body reddish-brown, apical 
third of elytra, apical abdominal segments and append- 
ages lighter, yellowish-brown. Pronotum and elytra 
not microsculptured. Antennae long, length/width ratio 


Scaphidinae of the Lesser Sunda Islands 95 


of antennomeres as follows: III 15/8: IV 37/6: V 47/7: 
VI 42/6: VII 48/12: VII 39/8: IX 50/12: X 47/13: XI 
58/14. Antennomere III subtriangular. Pronotum very 
finely punctate, punctures dense, not well delimited, 
lateral margins evenly rounded, lateral margin carinae 
not visible in dorsal view. Point of scutellum exposed. 
Elytra with lateral margin carinae visible near bases 
in dorsal view, apical margins truncate, inner apical 
angle rounded, not prominent, situated posterior level 
of outer angles, sutural margin slightly raised, sutural 
striae deep, weakly converging apically, not or hardly 
curved at base and not extending laterad pronotal lobe, 
adsutural areas flat. Elytral punctation fairly coarse, 
with punctures fairly well delimited, puncture intervals 
mostly about 1.5 to 2 times as large as puncture dia- 
meters. Hypomera smooth. Mesepimeron about as long 
as two thirds of interval to mesocoxa, about 4 times as 
long as wide. Metaventrite not microsculptured, flat in 
middle, with mesal groove and two fairly deep, elongate 
apicomedian impressions; punctation almost even, very 
fine and sparse. Submesocoxal areas 0.05 mm, about as 
two thirds of interval to metacoxa, submesocoxal lines 
convex, with fine marginal punctures. Metanepisterna 
flat, below level of metaventrite, narrowed anteriad, 
with inner margin straight, rounded at angles. Tibiae 
straight. Abdomen very finely punctate, with striate 
microsculpture. Sternite 1 with submetacoxal areas 
0.05-0.06 mm, as long as half of interval to apical mar- 
gin, submetacoxal lines strongly convex, with distinct 
margin punctures. 

Male: Tarsomeres 1 to 3 of protarsi slightly widened. 
Aedeagus (Figs 63-65) 0.36 mm long, asymmetrical, 
weakly sclerotized. Median lobe with basal bulb small, 
much shorter than apical process, articular process robust. 
Apical process narrow, strongly inflexed, parallel-sided 
and with blunt apex in dorsal view, tapering, with pointed 
apex, and most of ventral side oblique in lateral view. 
Parameres asymmetrical, almost straight, right paramere 
widest before middle, with shallow subapical notch, 
left paramere widest posterior mid-length, irregularly 
narrowed apically. Internal sac lacking flagellum or 
other sclerotized pieces, membranes lacking obvious 
denticulate or scale-like structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Senaro, N slope of Rinjani, 
1100 m. 


Comments: This species shares with S. ablutum the 
mesally grooved and apicomesally impressed metaven- 
trite. These two species resemble also by the col- 
our pattern of the elytra, and both have asymmetrical 
median lobe. They may be easily distinguished by the 
size of their body, significantly larger in S. ablutum. 
Scaphisoma adjunctum is characterized by the unique 
shape of the parameres. 


Scaphisoma aequum sp. nov. 
Fig. 66-67 


Holotype: SMNS; ©; Indonesia, W. Sumbawa 
Batudulang, 30 km S of Sumb. Besar, 1000 m Bolm Igt. 
10. Febr. 1994. 


Paratypes: SMNS, MHNG; 2 ¢, 
data as the holotype. 


4 ©; with the same 


Etymology: The species epithet is a Latin adjective, 
meaning same, similar. 


Description: Length 1.75-1.80 mm, width 1.12-1.15 
mm. Head and most of body uniformly dark red- 
dish-brown to black. Elytron with light reddish sub- 
apical band, narrow area posterior band usually 
reddish-brown, lighter than most of elytra and prono- 
tum. Abdomen, femora and tibiae almost evenly red- 
dish-brown, or sternite 1 darkened, almost as dark as 
metaventrite. Tarsi and antennae slightly lighter than 
tibiae. Pronotum and elytra lacking microsculpture. 
Antennae long, length/width ratio of antennomeres as 
follows: III 12/8: IV 40/6: V 50/7: VI 45/7: VII 53/10: 
VII 45/8: IX 52/13: X 50/13: XI 55/14. Antennomere 
III subtriangular. Pronotum very finely punctate, with 
lateral margins evenly rounded, lateral margin carinae 
not visible in dorsal view. Point of scutellum exposed. 
Elytra with lateral margin carinae entirely or partly 
exposed in dorsal view, apical margins truncate, inner 
apical angle rounded, not prominent, situated posterior 
level of outer angles, sutural margin not raised, sutural 
striae fairly deep, converging apically, at base hardly 
curved, not extending laterad pronotal lobe, adsutural 
areas flat. Elytral punctation sparse and fine near base, 
fairly coarse and dense on remaining surface. Hypomera 
smooth. Mesepimera about as long as interval to meso- 
coxa. Metaventrite very finely punctate, not microsculp- 
tured, with median part flattened, conspicuous mesal 
groove reaching anterior margin, and two small, shallow 
apicomedian impressions. Submesocoxal areas 0.06- 
0.07 mm, about as long as half of interval to metacoxa, 
submesocoxal lines convex, with very fine marginal 
punctures. Metanepisternum flat, narrowed apically, 
with inner margin oblique, impressed below margin 
of metaventrite. Tibiae straight. Abdomen very finely 
punctate, with microsculpture consisting of transverse 
striae. Submetacoxal areas 0.06-0.07 mm, submetacoxal 
lines convex, with distinct margin punctures. 

Male: Protarsal segments 1 to 3 hardly widened. 
Aedeagus (Figs 66-67) 0.37-0.40 mm long. Median lobe 
asymmetrical, weakly sclerotized. Basal bulb moderately 
large, articular process not prominent. Apical process 
almost in axis with basal lobe, weakly narrowed apically 
and with blunt tip in dorsal view, arcuate, abruptly 
narrowed posterior membranous fold and with tip acute 
in lateral view. Parameres asymmetrical, irregularly bent 
in dorsal view, with inner margin weakly sclerotized in 
apical thirds, almost evenly arcuate and narrow in lateral 


96 


I. Löbl 


NAME 
. N 


66 


Figs 62-69 
(62) Scaphisoma ablutum sp. nov., aedeagus in lateral view; scale bar = 0.2 mm. 
(63-65) Scaphisoma adjunctum sp. nov., aedeagus in dorsal (63) and lateral (64) views, parameres (65) in ventral view; scale bars = 


0.1 mm. 
(66-67) Scaphisoma aequum sp. nov., aedeagus in dorsal (66) and lateral (67) views. 
(68-69) Scaphisoma aereum sp. nov., aedeagus with extruded internal sac in dorsal view (68) and with internal sac in repos, in lateral 


(69) view; scale bars = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 97 


view. Internal sac with very narrow flagellum, surrounded 
by membranes. 


Distribution: Indonesia: Sumbawa. 


Type locality: W. Sumbawa, Batudulang, 30 km S of 
Sumb. Besar, 1000 m. 


Comments: The species shares many aedeagal charac- 
ters with S. adjunctum. It may be easily distinguished 
from the latter by the larger body size, colour and punc- 
tation of the elytra, and by the shape of the parameres. 


Scaphisoma aereum sp. nov. 
Figs 68-69 


Holotype: SMNS; 4; Indonesia, W. Sumbawa 
Batudulang, 30 km S of Sumb. Besar, 1000 m Bolm Igt. 
10. Febr. 1994. 


Paratypes: SMNS, MHNG; 2 <; with the same data as 
the holotype. 


Etymology: The species epithet is a Latin adjective, 
meaning sheathed. 


Description: Length 1.70-1.75 mm, width 1.07-1.10 
mm. Head and most of body uniformly blackish-brown 
to black, abdomen and femora reddish-brown, tibiae, 
antennae and tarsi lighter, almost yellowish. Pronotum 
and elytra lacking microsculpture, very finely punctate. 
Antennae long, length/width ratio of antennomeres as 
follows: III 12/7: IV 38/6: V 48/6: VI 45/7: VII 51/12: 
VIII 35/9: IX 53/13: X 40/15 (XI missing in all spe- 
cimens). Antennomere III short, subtriangular. Lateral 
margins of pronotum evenly rounded, lateral margin 
carinae not visible in dorsal view. Point of scutellum 
exposed. Elytra with lateral margin carinae exposed in 
dorsal view, apical margins truncate, inner apical angle 
rounded, not prominent, situated posterior level of outer 
angles, sutural margin not raised, sutural striae fairly 
shallow, parallel, at base somewhat curved, not extend- 
ing laterad pronotal lobe, adsutural areas flat. Elytral 
punctation sparse and very fine near base, becoming 
more dense and less fine apically, punctures distinctly 
larger than those on pronotum. Hypomera smooth. 
Mesepimera about as long as two thirds of interval 
to mesocoxa. Metaventrite not microsculptured, with 
median part flattened, conspicuous mesal groove and 
two foveiform apicomedian impressions. Submesocoxal 
areas 0.05 mm, about as long as fourth of interval to 
metacoxa, submesocoxal lines convex, with very fine 
marginal punctures. Metanepisternum flat, narrowed 
toward angles, with inner margin weekly rounded, 
impressed below margin of metaventrite. Tibiae straight. 
Abdomen with microsculpture consisting of transverse 
striae. Submetacoxal areas 0.07 mm, submetacoxal lines 
convex, with distinct margin punctures. 

Male: Protarsal segments | to 3 hardly widened. Aedea- 


gus (Figs 68-69) 0.38-0.41 mm long, fairly sclerotized. 
Median lobe asymmetrical, with small basal bulb, articu- 
lar process robust, not prominent, apical process abruptly 
curved and narrowed in dorsal view, arcuate and taper- 
ing in lateral view, tip acute in dorsal and lateral views. 
Parameres somewhat asymmetrical, almost evenly wide, 
weakly arcuate in lateral view, sinuate in dorsal view, 
inner margins not membranous, pores scattered in basal 
halves. Internal sac with moderately sclerotized flagel- 
lum and very finely denticulate membranes. 


Distribution: Indonesia: Sumbawa. 


Type locality: W. Sumbawa, Batudulang, 30 km S of 
Sumb. Besar, 1000 m. 


Comments: This species appears closely related with 
S. aequum. Both possess metaventrite with a median 
groove and a pair of foveiform impressions, and have 
similar aedeagi. These species may be easily distin- 
guished by their colour pattern. In addition, the groove 
of the metaventrite is shorter, the elytral punctation 
finer, and the apical process of the median lobe is 
abruptly narrowed in S. aereum. 


Scaphisoma adscitum sp. nov. 
Figs 70-71 


Holotype: SMNS; ©; Indonesia, Lombok Is. Senaro N 
slope of Rinjani 2.-5.Feb.1994 Bolm Igt. 1100 m. 


Paratype: MHNG; <; with the same data as the holo- 
type. 


Etymology: The species epithet is a Latin adjective, 
meaning strange. 


Description: Length 1.34-1.36 mm, width 0.90- 
0.92 mm. Head and most of body reddish-brown, apical 
margins of elytra and appendages lighter than body. 
Pronotum and elytra not microsculptured. Antennae 
short, length/width ratio of antennomeres as follows: 
III 6/7: IV 28/4: V 28/5: VI 31/7: VII 37/12: VII 
34/9: IX 40/12: X 37/13: XI 50/13. Antennomere III 
short, subtriangular. Pronotum very finely punctate, 
punctures sparse, not well delimited, lateral margins 
evenly rounded, lateral margin carinae not visible 
in dorsal view. Point of scutellum exposed. Elytra 
with lateral margin carinae hardly visible in dorsal 
view, apical margins truncate, inner apical angle 
rounded, not prominent, situated posterior level of 
outer angles, sutural margin not raised, sutural striae 
shallow, parallel, curved at base, not extending laterad 
pronotal lobe, adsutural areas flat. Elytral punctation 
very fine and dense near bases, distinct on remaining 
surface, with punctures fairly well delimited, puncture 
intervals mostly about 1.5 to 2.5 times as large as 
puncture diameters. Hypomera smooth. Mesepimeron 
about as long as interval to mesocoxa, about 4 times 


98 


as long as wide. Metaventrite not microsculptured, 
convex in middle, without impressions; punctation 
almost even, very fine and sparse. Submesocoxal 
areas 0.05 mm, about as two thirds of interval to 
metacoxa, submesocoxal lines convex, with fine 
marginal punctures. Metanepisterna flat, in level with 
metaventrite, narrowed anteriad, with inner margin 
straight, rounded at angles. Tibiae straight. Abdomen 
very finely punctate, with striate microsculpture 
becoming obsolete on lateral parts of sternite 1. 
Submetacoxal areas 0.06 mm, as long as half of interval 
to apical margin of sternite, submetacoxal lines strongly 
convex, With fine margin punctures. 

Male: Tarsomeres 1 to 3 of protarsi slightly widened. 
Aedeagus (Figs 70-71) 0.29-0.31 mm long, symmetrical, 
fairly sclerotized. Median lobe with basal bulb narrow, 
about as long as apical process, articular process not 
prominent. Apical process inflexed, weakly narrowed 
apically and with blunt apex in dorsal view, tapering, 
apex acute, ventral side arcuate in lateral view. Parameres 
wide near base and almost evenly broad in apical two 
third in dorsal view, narrowed to apical third in lateral 
view, not lobed, hardly bent in dorsal view. Internal sac 
without flagellum or other sclerotized pieces, membranes 
bearing denticulate structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Senaro N slope of Rinjani, 
1100 m. 


Comments: This species resembles S. antennarum 
described below but has much smaller submetacoxal 
areas. The aedeagal characters suggest relationship 
with S. /ombokense Löbl, 1986 though the aedeagus is 
smaller, the median lobe has prominent articular process 
and the apical process is comparatively much shorter. 
Scaphisoma lombokense differs in external characters 
notably by the sutural striae of the elytra extended along 
bases to form basal striae, and the antennomere V much 
longer than IV. 


Scaphisoma activum sp. nov. 
Figs 72-73 


Holotype: SMNS; €; Indonesia, W. Sumbawa 
Batudulang, 30 km S of Sumb. Besar, 1000 m Bolm Igt. 
10. Febr. 1994. 


Paratypes: SMNS, MHNG; 2 ¢, 2 ©; with the same 
data as the holotype 


Etymology: The species epithet is a Latin adjective, 
meaning active. 


Description: Length 1.45-1.60 mm, width 0.93-1.05 
mm. Most of body reddish-brown to blackish-brown, 
head, apices of elytra, apex of abdomen, femora and 
tibiae lighter, antennae and tarsi almost yellowish. 


Pronotum and elytra lacking microsculpture. Antennae 
moderately long, length/width ratio of antennomeres as 
follows: III 12/9: IV 32/8: V 32/9: VI 35/9: VII 38/13: 
VII 36/9: IX 39/12: X 42/12: XI 50/14. Antennomere 
III subtriangular. Pronotum very finely punctate, with 
lateral margins evenly rounded, lateral margin carinae 
not visible in dorsal view. Minute point of scutellum 
exposed. Elytra with lateral margin carinae exposed 
in dorsal view, apical margins truncate, inner apical 
angle rounded, not prominent, situated posterior level 
of outer angles, sutural margin not raised, sutural striae 
fairly shallow, parallel, at base somewhat curved, not 
extending laterad pronotal lobe, adsutural areas flat. 
Elytral punctation very fine near bases and along lat- 
eral margins, fairly coarse and dense on remaining dis- 
cal surface, with punctures well delimited and puncture 
intervals mostly about 1.5 to 2 times as large as punc- 
ture diameters. Hypomera smooth. Mesepimera some- 
what shorter than interval to mesocoxa. Metaventrite not 
microsculptured, with median area slightly convex, two 
very shallow apicomedian impression, punctation on 
apicomedian area fine but distinct, that on lateral areas 
very fine. Submesocoxal areas 0.05 mm, about as third 
of interval to metacoxa, submesocoxal lines convex, 
with very fine marginal punctures. Metanepisternum 
flat, narrowed anteriad, with inner margin rounded at 
angles, impressed below margin of metaventrite. Tibiae 
straight. Abdomen with microsculpture consisting of 
transverse striae, except on lateral parts of sternite 1 
lacking microsculpture. Submetacoxal areas 0.07-0.09 
mm, submetacoxal lines convex, with distinct margin 
punctures. 

Male: Protarsal segments 1 to 3 hardly widened. 
Aedeagus (Figs 72-73) 0.38-0.40 mm. Median lobe 
and parameres symmetrical. Median lobe moderately 
sclerotized, with basal bulb somewhat longer than apical 
process, articular process not prominent, apical process 
tapering, arcuate, with concave ventral side and acute 
tip in lateral view, weakly narrowed and with blunt tip 
in dorsal view. Parameres slightly curved and gradually 
narrowed in dorsal view, oblique, strongly narrowed from 
base to middle third and evenly wide in distal two thirds 
in lateral view; with scattered pores. Internal sac lacking 
sclerites, with very fine spinose structures in apical half, 
membranes with scale-like structures in basal half. 


Distribution: Indonesia: Sumbawa. 


Type locality: W. Sumbawa Batudulang, 30 km S of 
Sumb. Besar, 1000 m. 


Comments: The aedeagal characters suggest relation- 
ships with S. /ombokense Lôbl, 1986. This new spe- 
cies may be readily distinguished by the comparatively 
longer basal bulb of the aedeagus, the internal sac with a 
central tuft of denticles, the elytra with sutural striae not 
extended along basal margins, the comparatively much 
coarser elytral punctation, and the larger submesocoxal 
areas. 


Scaphidiinae of the Lesser Sunda Islands 99 


Scaphisoma acutatum Sp. nov. 
Figs 74-75 


Holotype: SMNS; ©; Indonesia, Lombok Is. Sapit- 
Sembalun Bumbung 14-16 Feb.1994 Bolm Igt., 900- 
1500 m. 


Paratype: MHNG; ©; with the same data as the holo- 
type. 


Etymology: The species epithet is a Latin adjective, 
meaning pointed and referring to the shape of the apex 
ofthe median lobe. 


Description: Length 1.85 mm, width 1.22 mm. Head, 
body and appendages almost evenly light brown, ti- 
biae and tarsi slightly lighter than pronotum or elytra. 
Antennae long, length/width ratio of antennomeres as 
follows: III 14/8: IV 16/7: V 37/9: VI 53/11: VII 58/13: 
VIII 48/8: IX 60/14: X 57/15: XI 64/12. Pronotum very 
finely punctate, punctures dense, not well delimited, 
lateral margins hardly rounded, lateral margin carinae 
not visible in dorsal view. Point of scutellum exposed. 
Elytra with lateral margin carinae exposed in dorsal 
view, apical margins rounded, inner apical angle not 
prominent, situated somewhat posterior level of outer 
angles, sutural margin not raised, sutural striae shallow, 
parallel, curved at base and extending laterally about 
to basal mid-width, adsutural areas flat. Elytral punc- 
tation very fine and dense, similar to pronotal anterior 
mid-length, posterior mid-length consisting of punctures 
larger and not well delimited, with puncture intervals 
mostly about twice to three times as large as puncture 
diameters. Hypomera smooth. Mesepimeron about 1.5 
times as interval to mesocoxa, about 4 times as long 
as wide. Metaventrite not microsculptured, slightly 
convex in middle, lacking impressions or sulci; punc- 
tation even, very fine and dense. Submesocoxal areas 
0.04 mm, about as fifth of interval to metacoxa, sub- 
mesocoxal lines convex, with fine marginal punctures. 
Metanepisternum flat, in level with metaventrite, not 
narrowed anteriad, with inner margin straight. Tibiae 
straight. Abdomen very finely punctate, with striate 
microsculpture. Submetacoxal areas 0.06 mm, sub- 
metacoxal lines convex, with fine margin punctures. 
Male: Tarsomeres 1 to 3 of protarsi slightly widened, 
Aedeagus (Figs 74-75) 0.58 mm long, symmetrical, fairly 
sclerotized. Median lobe with basal bulb narrow, longer 
than apical process, lacking prominent articular process. 
Apical process fairly inflexed, gradually narrowed api- 
cally in dorsal view, with ventral side arcuate in lateral 
view. Parameres narrow, not lobed, hardly bent and nar- 
rowed in subapical parts in dorsal view, almost straight 
and slightly widened apically in lateral view. Internal sac 
without flagellum or other sclerotized pieces, membranes 
bearing denticulate structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Sapit-Sembalun Bumbung, 
900-1500 m. 


Comments: The aedeagus of this species is similar to 
that of S. irideum Löbl, 2012 from the Moluccas island 
Halmahera, though the basal bulb is longer and the 
membranes of the internal sac are finely denticulate. It 
differs drastically from the latter by the not iridescent 
body, and from the congeners known from the Lesser 
Sundas by its short antennomere IV. 


Scaphisoma affabile sp. nov. 
Figs 76-77 


Holotype: MHNG; <; Indonesia, Indo: Timor (13) 
NTT: 16 km N Soe 30.3.91, evegrfor. D. Agosti, 
F91653. 


Etymology: The species epithet is a Latin adjective, 
meaning obliging, kindness. 


Description: Length 1.55 mm, width 1.07 mm. Head 
and most of body uniformly dark reddish-brown, apical 
parts of elytra light, yellowish, somewhat transparent. 
Light apical area of elytra extended from apical mar- 
gin to middle third near lateral margins, narrower near 
sutural striae. Appendages light reddish-brown. Sternite 
1 near base and apical sternites as dorsum of body, most 
of sternite | darkened. Antennae long, length/width ratio 
of antennomeres as follows: II 12/9: IV 35/6: V 40/7: 
VI 37/7: VII 45/10: VIII 32/10: IX 43/13: X 40/13: XI 
48/14. Antennomere III subtriangular. Pronotum and 
elytra lacking microsculpture, very finely punctate, 
punctures on elytra larger and deeper between mid- 
length and light apical area than on remaining elytral 
surface. Lateral margins of pronotum evenly rounded, 
lateral margin carinae not visible in dorsal view. Point 
of scutellum exposed. Elytra with lateral margin cari- 
nae not visible in dorsal view, apical margins weakly 
convex, inner apical angle rounded, not prominent, 
situated posterior level of outer angles, sutural margin 
not raised, sutural striae fairly deep, parallel, at base 
somewhat curved, not extending laterad pronotal lobe, 
adsutural areas flat. Hypomera smooth. Mesepimeron 
about as long as interval to mesocoxa. Metaventrite not 
microsculptured, lacking mesal impressions or sulci. 
Submesocoxal areas 0.05 mm, as long as half of interval 
to metacoxa, submesocoxal lines convex, with very fine 
marginal punctures. Metanepisternum flat, hardly nar- 
rowed toward angles, with inner margin almost straight 
and impressed below margin of metaventrite. Tibiae 
straight. Abdomen with microsculpture consisting of 
transverse striae. Submetacoxal areas 0.06 mm, sub- 
metacoxal lines convex, with short margin striae. 

Male: Protarsal segments 1 to 3 hardly widened. Aedea- 
gus (Figs 76-77) 0.53 mm long, moderately sclerotized. 
Median lobe weakly asymmetrical, with small basal 
bulb, articular process robust, not prominent, apical pro- 
cess strongly curved, tapering apically, tip acute in lateral 
view, blunt in dorsal view. Parameres symmetrical, wide- 


100 I. Löbl 


(os 


ul 
Ie N 
fis 
/ A 
1; i 
4 N 
na À 
1? NI 
ly \ 
11 \ 
\! \ 
x 


74 75 76 


Figs 70-77 

(70-71) Scaphisoma adscitum sp. nov., aedeagus in dorsal (70) and lateral (71) views. 

(72-73) Scaphisoma activum sp. nov., aedeagus in dorsal (72) and lateral (73) views. 

(74-75) Scaphisoma acutatum sp. nov., aedeagus in dorsal (74) and lateral (75) views. 

(76-77) Scaphisoma affabile sp. nov., aedeagus in dorsal (76) and lateral (77) views; scale bars = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 101 


ned in apical halves, inner margins membranous, pores 
scattered in proximal halves of parameres. Internal sac 
with moderately sclerotized flagellum, apical parts of 
membranes with very fine denticulate structures. 


Distribution: Indonesia: Timor. 
Type locality: Timor, 16 km N Soe. 


Comments: The aedeagal characters suggest rela- 
tionships with the New Guinean S. pseudofasciatum 
Löbl, 1975 and S. triste Löbl. 1975, though S. pseudo- 
fasciatum possesses apical process of the median lobe 
abruptly narrowed and internal sac extruded in repos. 
Both species may be easily distinguished by their body 
colour. Scaphisoma triste is larger, its body is over 
2.1 mm long and has elytra with sutural striae extended 
along bases to form basal striae; it differs also conspicu- 
ously by the strongly prominent articular process of the 
median lobe. 


Scaphisoma antennarum sp. nov. 
Figs 78-79 


Holotype: SMNS; <@; Indonesia, Lombok Is. Sapit- 
Sembalun Bumbung 14-16 Feb.1994 Bolm Igt. 900- 
1500 m. 


Paratypes: SMNS, MHNG; 6 d, 1 ©; with the 
same data as the holotype. - SMNS; 1 ©; Indonesia 
E. Lombok Sapit, 14.-16.2.1994 SE slope of Mt. Rinjani 
Bolm let. 


Etymology: The species epithet is a Latin adjective, 
referring to the antennae. 


Description: Length 1.35-1.50 mm, width 0.96-1.06 
mm. Head and body dark reddish-brown to blackish- 
brown. Elytra eventually darkened apically, except for 
narrowly light apices. Apical abdominal segments and 
appendages light reddish to yellowish. Pronotum and 
elytra lacking microsculpture. Antennae short, length/ 
width ratio of antennomeres as follows: III 10/10: IV 
15/8: V 22/9: VI 27/12: VII 42/14: VII 26/11: IX 40/12: 
X 38/16: XI 50/16. Antennomere III subtriangular. 
Pronotum very finely punctate, lateral margins evenly 
rounded, lateral margin carinae not visible in dorsal 
view. Point of scutellum exposed. Elytra with lateral 
margin carinae visible in dorsal view, apical margins 
weakly convex, inner apical angle rounded, not 
prominent, situated posterior level of outer angles, 
sutural margin not raised, sutural striae fairly shallow, 
parallel, somewhat curved at base, not extending laterad 
pronotal lobe, adsutural areas flat. Punctation very fine 
and sparse on basal fourth to third of elytra, becoming 
less fine and denser apically, punctures on apical halves 
of elytral disc well delimited, to part about as large as 
puncture intervals. Hypomera smooth. Mesepimeron 
about as long as interval to mesocoxa. Metaventrite 
not microsculptured, lacking mesal impressions or 


sulci, punctation very dense and fine on apicomedian 
area, with punctures to part as large as intervals, 
finer and sparser on anteriomedian and lateral areas. 
Submesocoxal areas 0.04-0.05 mm, as long as fourth 
of interval to metacoxa, submesocoxal lines convex, 
with very fine marginal punctures. Metanepisternum 
flat, slightly narrowed toward angles, with inner 
margin weakly sinuate, impressed below margin of 
metaventrite. Tibiae straight. Abdomen lacking obvious 
microsculpture, pygidium conspicuously punctured, 
with punctures dense, about as large as puncture 
intervals, sternites very finely and sparsely punctured, 
excepted on median part of sternite 1 with denser 
punctures, similar to those on apicomedian part of 
metaventrite. Submetacoxal areas 0.12-0.13 mm, about 
twice as shortest interval to apical margin of sternite, 
submetacoxal lines strongly convex, with fine margin 
punctures. 

Male: Protarsal segments | to 3 hardly widened. Aedea- 
gus (Figs 78-79) 0.23-0.27 mm long. Median lobe 
strongly sclerotized, slightly asymmetrical, with basal 
bulb small, bearing comparatively large articular process. 
Apical process much longer than basal bulb, narrow, 
tapering, in dorsal view weakly curved, in lateral view 
arcuate basally, straight in apical half. Parameres fairly 
narrow, dorsally widened at apices. Internal sac with 
hardly visible flagellum, membranes lacking denticulate 
or scale-like structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Sapit-Sembalun Bumbung, 
900-1500 m. 


Comments: The aedeagus in the new species is similar 
to that in S. besucheti Löbl, 1971 from Sri Lanka, 
though it is significantly smaller. These two species 
may be readily distinguished also by their antennae, 
the elytra lacking basal striae in S. antennarum, and the 
much smaller submetacoxal areas in S. besucheti. 


Scaphisoma animatum sp. nov. 
Figs 80-81 


Holotype: SMNS; ©; Indonesia, Lombok Is. Sapit- 
Sembalun Bumbung 14.-16.Feb.1994 Bolm Igt. 900- 
1500 m. 


Paratypes: SMNS, MHNG; | ©, 2 ©; with the same 
data as the holotype. 


Etymology: The species epithet is a Latin adjective, 
meaning courageous. 


Description: Length 1.37-1.46 mm, width 0.94- 
0.96 mm. Head and most of body dark brown or red- 
dish-brown. Most of elytra light, yellowish, basal fourth 
of elytra about as pronotum or somewhat darker, ante- 
rior halves of sutural areas slightly darkened, and each 
elytron with darkened subapical band. Ventral side of 


102 I. Löbl 


4 
À 
À 
i 
À 
N 
a 
x 
N 
v 


Figs 78-84 
(78-79) Scaphisoma antennarum sp. nov., aedeagus in dorsal (78) and lateral (79) views. 
(80-81) Scaphisoma animatum sp. nov., aedeagus in dorsal (80) and lateral (81) views. 


(82-84) Scaphisoma acutum sp. nov., aedeagus in dorsal (82) and lateral (83) views, apical part of median lobe with basal parts of 
parameres (84) in dorsal view; scale bars = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 103 


thorax about as dark as pronotum. Apical abdominal 
segments and appendages light, yellowish. Pronotum 
and elytra lacking microsculpture. Antennae long, 
length/width ratio of antennomeres as follows: III 12/7: 
IV 20/5: V 33/6: VI 40/8: VII 49/13: VII 40/10: IX 
48/12: X 45/11: XI 54/14. Antennomere III short, sub- 
triangular. Pronotum very finely punctate, lateral mar- 
gins evenly rounded, lateral margin carinae not visible 
in dorsal view. Point of scutellum exposed. Elytra with 
lateral margin carinae visible in dorsal view, apical 
margins weakly convex, inner apical angle rounded, 
not prominent, situated posterior level of outer angles, 
sutural margin not raised, sutural striae fairly shallow, 
parallel, somewhat curved at base, not extending late- 
rad pronotal lobe, adsutural areas flat. Elytral puncta- 
tion very fine and sparse, similar to pronotal near bases, 
more distinct on most of discal surface and with punc- 
ture intervals about 2 to 3 times as large as puncture 
diameters. Hypomera smooth. Mesepimeron slightly 
longer than interval to mesocoxa. Metaventrite not 
microsculptured, lacking mesal impressions or sulci, 
very finely punctate. Submesocoxal areas 0.04 mm, 
as long as third of interval to metacoxa, submeso- 
coxal lines convex, with fine marginal punctures. 
Metanepisternum flat, slightly narrowed toward angles, 
with middle part of inner margin oblique, not impressed 
below margin of metaventrite. Tibiae straight. Abdomen 
very finely punctate, sternite 1 lacking microsculp- 
ture, apical sternites with punctate microsculpture. 
Submetacoxal areas 0.07-0.09 mm, almost reaching 
sternal mid-length, submetacoxal lines convex, with 
coarse margin punctures. 

Male: Protarsal segments 1 to 3 hardly widened. Aedea- 
gus (Figs 80-81) 0.25 mm long, moderately sclero- 
tized. Median lobe symmetrical, with large basal bulb, 
articular process robust and prominent, apical process 
much shorter than basal bulb, strongly curved, tapering 
apically, tip acute in lateral view, blunt in dorsal view. 
Parameres symmetrical, narrow, almost straight in dorsal 
view, hardly bent and widened at apices in lateral view. 
Internal sac lacking flagellum and sclerotized pieces, 
membranes with fine denticulate structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Sapit-Sembalun Bumbung, 
900-1500 m. 


Comments: The aedeagal characters suggest 
relationship with S. /ombokense Löbl, 1986 and S. bugi 
Löbl, 1983, although these two species have parameres 
widened basally, and may be easily distinguished by 
their colour pattern and abdominal microsculpture 
(abdomen with striate microsculpture, body uniformly 
reddish-brown in S. /ombokense, elytra dark with lighter 
reddish subbasal and subapical spots in S. bugi; see 
Löbl, 1983 and 1986, respectively). See also comments 
under S. ascitum. 


Scaphisoma acutum sp. nov. 
Figs 82-84 


Holotype: MHNG; <; Indonesia, Indo: Lombok 
Batu Koq (N of G. Rinjani) 500 m, sec.forest in gorge 
D. Agosti 12.03.1991. 


Etymology: The species epithet is a Latin adjective, 
meaning pointed and referring to pointed sclerites of the 
internal sac. 


Description: Length 1.22 mm, width 0.82 mm. Head 
and body dark reddish-brown, appendages light. 
Pronotum and elytra lacking microsculpture. Antennae 
long, length/width ratio of antennomeres as follows: III 
12/6: IV 28/5: V 40/6: VI 35/7: VII 38/11: VII 37/8: IX 
42/12 (left antennomeres X and XI, and right antenno- 
mere VIII-XI broken off). Antennomere III subtriangu- 
lar. Pronotum finely punctate, punctures on basomedian 
area larger than near lateral and apical margins, lateral 
margins evenly rounded, lateral margin carinae visible 
in dorsal view. Point of scutellum exposed. Elytra with 
lateral margin carinae visible in dorsal view, apical mar- 
gins truncate, inner apical angle rounded, not prominent, 
situated posterior level of outer angles, sutural margin 
not raised, sutural striae fairly deep, converging api- 
cally, somewhat curved at base, not extending laterad 
pronotal lobe, adsutural areas flat. Elytral punctation 
coarse and dense, punctures fairly well delimited, punc- 
ture intervals mostly about as large to twice as large as 
puncture diameters. Hypomera smooth. Mesepimeron 
about twice as interval to mesocoxa, about 5 times as 
long as wide. Metaventrite not microsculptured, lack- 
ing mesal impressions or sulci, punctation even, very 
fine and sparse. Submesocoxal areas 0.05 mm, almost as 
long as half of interval to metacoxa, submesocoxal lines 
convex, with fine marginal punctures. Metanepisternum 
flat, impressed below margin of metaventrite, slightly 
narrowed anteriad, with inner margin weakly sinuate. 
Tibiae straight. Abdomen very finely punctate. Sternite 
1 with strigulate microsculpture distinct in middle, 
becoming punctulate or obsolete laterally, with not 
well delimited lateral impression. Submetacoxal areas 
0.05 mm, submetacoxal lines convex, with fine margin 
punctures. 

Male: Protarsal segments | to 3 hardly widened. Aedea- 
gus (Figs 82-84) 0.37 mm long, weakly sclerotized, sym- 
metrical. Median lobe with basal bulb much longer than 
apical process, lacking prominent articular process. Api- 
cal process strongly inflexed and narrowed apically, con- 
cealed in dorsal view. Parameres wide in basal halves and 
lobed, bent in middle, narrow in apical halves in dorsal 
view, gradually narrowed apically in lateral view. Inter- 
nal sac lacking robust sclerites, with short denticulate 
structures in proximal part, long, very narrow and acute 
denticles in apical part. 


Distribution: Indonesia: Lombok. 


104 


I. Lobl 


SZ 


87 


1990. 


Dt 
170877: 


89 


Figs 85-90 
(85-86) Scaphisoma aproximatum sp. nov., aedeagus in dorsal (85) and lateral (86) views; scale bar = 0.1 mm. 
(87-90) Scaphisoma angulare sp. nov., aedeagus in dorsal (87) view, apical part of median lobe with parameres (88) in dorsal view, 


basal half of internal sac (89) and apical half of internal sac (90) in dorsal view; scale bars = 0.2 mm. 


Scaphidiinae of the Lesser Sunda Islands 105 


Type locality: Lombok, Batu Koq (N of G. Rinjani) 
500 m. 


Comments: The aedeagal characters suggest 
relationships of S. acutum with S. sesaotense Löbl, 
1986 and the Sumatran S. irregulare Löbl, 1975. The 
new species may be distinguished from S. sesaotense 
by the much smaller aedeagus with short parameral 
lobes and internal sac lacking large bifid sclerite. 
Scaphisoma irregulare has distinctive elytral punctation, 
with punctures on middle of disc coarser and also 
much denser than near margins. Unlike in S. acutum 
and S. sesaotense, S. irregular has parameres finely 
tuberculate, the parameral lobes very narrow, and does 
not possess sclerotized pieces in the internal sac. 


Scaphisoma approximatum sp. nov. 
Figs 85-86 


Holotype: MHNG; €; Indonesia, Indo.: Lombok Mt. 
Rinjani above Senaro, 900-1100 m 6. XI 1991, I. Löbl 
forest floor litter. 


Etymology: The species epithet is a Latin adjective, 
meaning being close. 


Description: Length 1.65 mm, width 1.14 mm. Head, 
body and appendages light reddish-brown. Pronotum 
and elytra lacking microsculpture. Antennae long, 
length/width ration of antennomeres as follows: III 
15/8: IV 29/6: V 35/6: VI 36/7: VII 48/15: VIII 35/9: 
IX 48/15: X 44/15: XI 65/15. Antennomere III subtri- 
angular. Pronotum very finely punctate, lateral mar- 
gins evenly rounded, lateral margin carinae not visible 
in dorsal view. Point of scutellum exposed. Elytra with 
lateral margin carinae visible in dorsal view, apical 
margins weakly convex, inner apical angle rounded, 
not prominent, situated posterior level of outer angles, 
sutural margin not raised, sutural striae fairly shallow, 
parallel, at base somewhat curved, not extending late- 
rad pronotal lobe, adsutural areas flat. Punctation very 
fine and sparse near elytral bases, becoming more dis- 
tinct and denser apically, punctures on apical halves of 
elytral disc well delimited, puncture intervals mostly 
about as large to twice as large as puncture intervals. 
Pygidium distinctly, densely punctate, most punctures 
about as large as puncture intervals. Hypomera smooth. 
Mesepimeron somewhat shorter than interval to meso- 
coxa, about 2.5 times as long as wide. Metaventrite 
not microsculptured, lacking mesal impressions or 
sulci, with punctation even, very fine and sparse. 
Submesocoxal areas 0.03 mm, as long as sixth of inter- 
val to metacoxa, submesocoxal lines parallel, with 
very fine marginal punctures. Metanepisternum flat, 
impressed below margin of metaventrite, narrowed ante- 
riad, with inner margin weakly sinuate. Tibiae straight. 
Abdomen lacking obvious microsculpture. Sternite | 
with punctation dense and comparatively coarse on 


basal half, becoming finer and sparser toward api- 
cal margin. Following sternites very finely punctate. 
Submetacoxal areas 0.02-0.03 mm, submetacoxal lines 
parallel, with fine margin punctures. 

Male: Protarsal segments 1 to 3 hardly widened. Aedea- 
gus (Figs 85-86) 0.37 mm long, weakly sclerotized, sym- 
metrical. Median lobe with basal bulb longer than apical 
process and large, prominent articular process. Apical 
process moderately narrowed apically in dorsal view, 
strongly inflexed, with oblique ventral side, abruptly 
narrowed in apical part in lateral view. Parameres wide, 
arcuate in dorsal view, strongly widened ventrally and 
abruptly narrowed at tip in lateral view. Internal sac with 
robust bidentate sclerite and finely denticulate mem- 
branes. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok Mt. Rinjani above Senaro, 900- 
1100 m. 


Comments: The species resembles S. incomptum Löbl, 
1986 from Sumatra, though it has conspicuously longer 
antennae. Its aedeagal characters suggest relationships 
with the Sumatran S. heissi Löbl, 1982, that possesses 
larger and strongly sclerotized aedeagus with the apical 
process of the median lobe parallel and evenly curved. 
The new species may be readily distinguished from 
S. heissi by the uniformly light body colour and the 
elytra with parallel sutural striae. 


Scaphisoma angulare sp. nov. 
Figs 87-90 


Holotype: SMNS; €; Indonesia, W. Sumbawa 
Batudulang, 30 km S of Sumb. Besar, 1000 m Bolm Igt. 
10. Febr. 1994. 


Paratypes: SMNS, MHNG; 7 ©, 1 ©; Indonesia, with 
the same data as the holotype. 


Etymology: The species epithet is a Latin adjective, 
meaning angulate and referring to the shape of the 
elytra. 


Description: Length 1.93-2.10 mm, width 1.25-1.33 
mm. Head, pronotum and hypomera very dark red- 
dish-brown to blackish-brown. Elytra black, with 
base somewhat lighter, adsutural areas about as pro- 
notum, at apices yellowish. Metaventrite very dark 
reddish-brown to blackish, abdomen dark brown or 
reddish-brown. Appendages light reddish-brown to 
yellowish. Pronotum and elytra lacking microsculp- 
ture. Antennae long, length/width ratio of antenno- 
meres as follows: III 16/8: IV 40/8: V 55/10: VI 51/9: 
VII 58/14: VII 51/10: IX 57/10: X 55/14: XI 65/15. 
Antennomere III subtriangular. Pronotum finely and 
densely punctate, with lateral margins evenly rounded, 
basal halves of lateral margin carinae visible in dorsal 
view. Apex of scutellum exposed. Elytra with lateral 


106 I. Lobl 


Figs 91-94 
(91-94) Scaphisoma affectum sp. nov., aedeagus in dorsal view (91), median lobe with parameres, without proximal part of basal bulb 
(92) in dorsal and (93) lateral views, interval sac (94) in dorsal view; scale bars = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 107 


margin carinae entirely exposed in dorsal view, apical 
margins truncate, inner apical angle rectangular, not 
prominent in male, prominent and tooth-like in female; 
situated posterior level of outer angles, sutural margin 
not raised, sutural striae deep, parallel, hardly curved 
at base, not extending laterad pronotal lobe, adsutural 
areas flat. Elytral punctation fairly coarse, dense, punc- 
tures well delimited, puncture intervals mostly about as 
large to twice as large as puncture diameters. Hypomera 
smooth. Mesepimera longer than interval to mesocoxa. 
Metaventrite with microsculpture striagulate, distinct 
on median area, hardly visible on lateral areas. Median 
area of metaventrite slightly convex, with two indis- 
tinct apicomedian impressions, punctation fine but dis- 
tinct apicomedianly, elsewhere very fine. Submesocoxal 
areas 0.04 mm, about as fifth of intervals to metacoxae, 
submesocoxal lines subparallel, with coarse marginal 
punctures. Metanepisternum flat, hardly narrowed ante- 
riad, with inner margin straight in middle, rounded near 
angles, impressed below margin of metaventrite. Tibiae 
straight. Abdomen with microsculpture consisting of 
transverse striae. Sternite 1 all over very finely punctate. 
Submetacoxal areas 0.07-0.08 mm, submetacoxal lines 
convex, with coarse margin punctures. 

Male: Protarsal and mesotarsal segments | to 3 distinctly 
widened. Inner apical angle of elytra not prominent. 
Aedeagus (Figs 87-90) 1.07-1.10 mm long. Median lobe 
and parameres symmetrical, strongly sclerotized. Median 
lobe with large basal bulb, ventral branch of apical pro- 
cess short, weakly inflexed in apical half, with subapical 
dorsal denticle, dorsal branches of apical process slightly 
shorter than ventral branch, articular process not pro- 
minent. Parameres in level with median lobe, arcuate in 
basal part, oblique posterior basal part, narrowed toward 
apices in lateral view, sinuate in dorsal view, lacking 
distinct crenulations; pores scattered, situated in basal 
halves. Internal sac complex, with tufts of dense denti- 
cles, large median tooth, and membranes bearing scale- 
like structure. 

Female: Inner apical angle of elytra prominent, tooth- 
like. 


Distribution: Indonesia: Sumbawa. 


Type locality: W. Sumbawa Batudulang, 30 km S of 
Sumb. Besar, 1000 m. 


Comments: The aedeagus of this species is very simi- 
lar to that of S. sapitense, suggesting close relation- 
ships. Possibly, these species are vicariant. The new 
species differs however conspicuously from S. sapitense 
by its colour pattern, and also by the very finely punc- 
tate basomedian part of the sternite 1 and the laterally 
microsculptured metaventrite. The aedeagi are in both 
species similar. Unlike S. sapitense, S. angulare has 
parameres distinctly sinuate in dorsal view and its inter- 
nal sac bears a large median tooth, while it lacks the row 
of scale-like, flat sclerites. 


Scaphisoma affectum sp. nov. 
Figs 91-94 


Holotype: MHNG; €; Indonesia, Indo: Bali Mt. 
Batukaru 500-700 m, 1991 I. Löbl, 28.-29.X. 


Etymology: The species epithet is a Latin adjective, 
referring to the affected characters. 


Description: Length 1.85 mm, width 1.33 mm. Head, 
pronotum and most of elytra reddish-brown, api- 
cal fourth of elytra lighter, yellowish. Hypomera 
light brown. Mesoventrite, mesepisterna, metaven- 
trite, metanepisterna, metepimeres and sternite | dark 
brown. Remaining exposed abdominal segments lighter, 
appendages yellowish. Pronotum and elytra lacking 
microsculpture. Antennae long, length/width ratio of 
antennomeres as follows: III 12/8: IV 30/6: V 52/7: 
VI 55/8: VII 57/13: VII 45/10: IX 57/12: X 57/12: XI 
65/13. Antennomere III triangular. Pronotum finely and 
densely punctate, with lateral margins evenly rounded, 
lateral margin carinae hardly visible in dorsal view. 
Apex of scutellum exposed. Elytra with lateral margin 
carinae entirely exposed in dorsal view, apical margins 
rounded, inner apical angle situated in level with outer 
angles, sutural margin not raised; sutural striae fairly 
shallow, oblique near base and not extending late- 
rad pronotal lobe, almost parallel from oblique section 
to apices, adsutural areas flat. Elytral punctation fairly 
coarse, dense, punctures well delimited, puncture inter- 
vals mostly about as large to twice as large as punc- 
ture diameters. Hypomera finely punctate. Mesepimera 
about four times as long as wide, about as long as 
three fourth of interval to mesocoxa. Metaventrite with 
microsculpture between mesocoxae and metacoxae. 
Median area of metaventrite convex, without impres- 
sions or grooves, flattened near metacoxal process, with 
fairly coarse puncture rows parallel to metacoxae, punc- 
tation elsewhere very fine and sparse. Submesocoxal 
areas 0.05 mm, about as fourth of intervals to 
metacoxae, submesocoxal lines convex, with fairly 
indistinct marginal punctures. Metanepisternum convex, 
distinctly narrowed anteriad, with inner margin slightly 
impressed and almost straight except at rounded poste- 
rior angles. Mesotibiae and metatibiae slightly curved. 
Abdomen with microsculpture consisting of transverse 
striae. Sternite 1 with coarse puncture on narrow baso- 
median area, very finely punctate on remaining surface. 
Submetacoxal areas 0.05 mm, submetacoxal lines con- 
vex, with coarse margin punctures. 

Male: Protarsi with segment 1 distinctly widened, nar- 
rower than tibia, segments 2 and 3 slightly widened. 
Aedeagus (Figs 91-94) 1.0 mm long. Median lobe with 
large, moderately sclerotized basal bulb, arcticular 
process not projecting. Apical process asymmetrical, 
strongly inflexed, sinuate and tapering in lateral view, 
oblique in dorsal view. Parameres symmetrical, strongly 
expanded ventrally to form large, strongly sclerotized 


108 I. Löbl 


subbasal apophyses posterior bases. Dorsal margins of 
parameres strongly sclerotized posterior level of apophy- 
ses, wide areas below dorsal margins membranous. Inter- 
nal sac complex, with three strongly sclerotized apical 
teeth and basal plates, and membranes with scale-like 
and denticulate structures. 


Distribution: Indonesia: Bali. 
Type locality: Bali, Mt. Batukaru 500-700 m. 


Comments: This species is possibly related with S. rufi- 
colle (Pic, 1915) described from the East Malaysian 
island Bangii. Unfortunately, the apical process of the 
median lobe is hardly visible in the sole male of the 
latter species that is available for study. Both species 
clearly differ also by the parameres, gradually wid- 
ened and with postbasal lobes situated more distally 
in S. ruficolle, and by the structures of the internal sac, 
in particular by the presence of a single robust tooth in 
S. ruficolle. 


Scaphisoma aspectum sp. nov. 
Figs 95-98 


Holotype: SMNS; ; Indonesia, Bali Danau Buyan, 
1300 19.-21.2. 1994 Bolm let. 


Paratypes: SMNS, MHNG; 17 &, 9 ©; with the same 
data as the holotype. - SMNS, MHNG; 14 &, 14 9; 
Bali 12m km NW of Bedugul (Buyan Lake) 29 Apr.-2. 
May 2001 Bolm Igt., 950 m. 


Etymology: The species epithet is a Latin adjective, 
meaning aspect. 


Description: Length 1.95-2.15 mm, width 1.35- 
1.48 mm. Colour of head, pronotum and abdomen: see 
under male and female characters. Elytra dark brown 
to black, each with light, yellowish-brown or reddish 
subhumeral spot and subapical band, latter usually 
not clearly delimited. Venter of thorax, hypomera and 
mesoventrite. excepted, and sternite 1 dark brown to 
black, mesoventrite somewhat lighter, exposed tergites 
and appendages light, yellowish-brown. Pronotum and 
elytra of lacking microsculpture. Antennae long, length/ 
width ratio of antennomeres as follows: III 26/6: IV 
37/7: V 65/7: VI 63/9: VII 65/15: VIII 57/8: IX 72/12: 
X 62/12: XI 77/13. Antennomere III conspicuously 
elongate, subcylindrical. Pronotum finely and densely 
punctate, with lateral margins evenly rounded, lateral 
margin carinae usually visible in dorsal view. Apex of 
scutellum exposed. Elytra with lateral margin carinae 
entirely exposed in dorsal view, apical margins trun- 
cate, inner apical angle situated posterior level of outer 
angles, sutural margin slightly raised in posterior two 
thirds of sutural length, sutural striae deep, not curved 
at base and not extending laterad pronotal lobe, conspi- 
cuously converging apically, adsutural areas flat, each 


about 0.20 mm wide at level of scutellum and about 
0.10 mm wide at mid-length. Elytral punctation fairly 
coarse, dense, punctures well delimited, puncture inter- 
vals mostly about as large to twice as large as puncture 
diameters. Hypomera smooth. Mesepimera about 3 to 4 
times as long as wide, shorter than interval to mesocoxa. 
Metaventrite with striate microsculpture. Median area 
of metaventrite slightly convex, without impressions 
or grooves, with fairly coarse puncture rows parallel to 
metacoxae, punctation very fine and sparse elsewhere. 
Submesocoxal areas 0.04 mm, about as fifth of inter- 
vals to metacoxae, submesocoxal lines subparallel, 
with fairly coarse marginal punctures. Metanepisternum 
convex, weakly narrowed anteriad, with inner margin 
deeply impressed and almost straight. Mesotibiae and 
metatibiae slightly curved. Abdomen with microsculp- 
ture consisting of transverse striae. Sternite 1 all over 
very finely punctate. Submetacoxal areas 0.04 mm, sub- 
metacoxal lines convex, with fairly coarse margin punc- 
tures. 

Male: Head narrowly dark brown to black on vertex, 
light reddish-brown to yellowish-brown on remaining 
surface. Pronotum laterally and hypomera entirely light 
reddish-brown to yellowish-brown. Mesal part of pro- 
notum dark brown to black. Sternite 1 dark, about as 
mesoventrite, following ventrites lighter. Protarsal and 
mesotarsal segments | to 3 distinctly widened. Inner api- 
cal angle of elytra not prominent. Aedeagus (Figs 95-98) 
1.15-1.28 mm long. Median lobe and parameres asym- 
metrical. Median lobe with large, moderately sclerotized 
basal bulb, apical process strongly sclerotized, irregularly 
curved and inflexed, in lateral view wider than in dorsal 
view. Parameres each with large ventral lobe. Internal sac 
with two sclerites, left sclerite long and abruptly truncate 
at apex, right sclerite strongly widened in apical half; 
membranes bearing scale-like structures. 

Female: Head, pronotum and hypomera uniformly 
or almost uniformly dark reddish-brown to black, or 
head somewhat lighter than pronotum. Sternites, apical 
excepted, about as dark as metaventrite. 


Distribution: Indonesia: Bali. 


Comments: This species is closely related with and 
similar to S. testaceomaculatum (Pic, 1915) from Java. 
The males may be easily distinguished by the pronotal 
colour pattern. While the dark area in S. festaceomacu- 
latum is limited to a narrow band along the basal mar- 
gin of the pronotum and is just moderately extended 
mesally, it covers the entire mesal area in S. aspectum. 
The shape of the apical process of the median lobe and 
of the structures of the internal sac differ drastically in 
these two species. 

Achard (1920) downgraded Pseudoscaphosoma sub- 
elongatum Pic, 1915 to variety of P. testaceomacula- 
tum Pic, 1915. The study of the respective type material 
confirms the synonymy of these names. The “thorace 
posticis breve nigro notato” given by Pic for P. testaceo- 


Scaphidiinae of the Lesser Sunda Islands 109 


maculatum is a male character, absent in the description 
of P subelongatum. Both names are here fixed: 

P. testaceomaculatum Pic, lectotype MNHN; d; by pres- 
ent designation, labelled: Java occident. Pengalengan 
4000’ 1893 H.Fruhstorfer / Type (handwritten)/ TYPE 
(red, printed) Pseudoscaphosoma testaceomaculatum Pic 
(handwritten by Pic) /Scaphisoma testaceomaculatum 
(Pic) der Löbl 1974 / /Lectotype Scaphisoma testaceo- 
maculatum (Pic) det. Löbl, 2014. 

P. subelongatum Pic, lectotype MNHN; ©; by present 
designation, labelled: Java occident. Pengalengan 4000’ 
1893 H.Fruhstorfer / Type (handwritten) / TYPE (red, 
printed) /? Toxidium Lec (handwritten by Pic)/ Pseu- 
doscaphosoma subelongatum Pic (handwritten by Pic) / 
Scaphisoma testaceomaculatum (Pic) der Löbl 1974 / / 
Lectotype Scaphisoma subelongatum (Pic) det. Löbl, 
2014. 


Scaphisoma coarctatum Löbl, 1976 


Additional material examined: SMNS; 1 4; 
Indonesia, Lombok, Sapit - Sembalun Bumbung, 14.- 
16.Feb 1994, Bolm Igt., 900-1400 m. 


Distribution: Indonesia: Buru, Lombok. 


Comments: The description of this species was based 
on a single male form Buru (in Zoölogisch Museum, 
Amsterdam, at present in NBCL). The specimen from 
Lombok has elytra with sutural striae longer, starting at 
elytral bases, at each side of the pronotal lobe. 


Scaphisoma dohertyi (Pic, 1915) 


Additional material examined: SMNS, MHNG; 
Indonesia, Bali, Danau Buyan, 1300 m, 19.-21.2.1994, 
Bolm Igt., 3 ex. — SMNS, MHNG; 11 ex; NE 
Sumbawa, Calabai (Tambora N.P.) 11.-13.Febr. 1994 
Bolm Igt. 


Distribution: Tropical Asia from north-eastern India to 
Yunnan, and Indonesia: Java, Bali and Sumbawa. 


Comments: The aedeagus of the species is illustrated in 
Löbl, 1981a. 


Scaphisoma gracilicorne Achard, 1920 
Figs 99-102 


Lectotype: MNHN; 4; by present designation, labelled: 
Sumatra Coll Grovelle (handwritten) TYPE (printed, 
red) / Scaphosoma gracilicorne TYPE (handwritten by 
Achard) J. Achard det. /Scaphisoma gracilicorne Achard 
det. Lòbl 1974 / Lectotype Scaphisoma gracilicorne 
Achard det. I. Löbl, 2014. The right elytron and the 
antennae from 3rd segment on are broken off and miss- 
ing. 


Additional material examined: SMNS; 1 4; 


Indonesia, Lombok, Senaro N slope of Rinjani, 2.-5. 


Feb.1994, 1100 m, Bolm Igt. - MNHN; 1 4; Sumatra, 
Fort de Kock [=Bukittinggi] 920 m 1925 leg. E. 
Jacobson. 


Redescription: Length 1.50-1.55 mm, width 1.0- 
1.05 mm. Head and body light reddish-brown to yel- 
lowish, middle part of elytra somewhat darkened, apical 
third to two fifth of elytra lighter. Apex of abdomen and 
appendages yellowish. Antennae fairly long, length/ 
width ratio of antennomeres as follows: III 12/7: IV 
25/6: V 36/6: VI 32/6: VII 40/10: VIII 36/6: IX 44/10: 
X 40/12: XI 50/12. Antennomere III subtriangular. 
Pronotum finely punctate, with lateral margins evenly 
rounded, lateral margin carinae hardly visible near 
base or in basal half in dorsal view. Apex of scutellum 
exposed. Elytra with lateral margin carinae visible near 
bases or in basal third, in dorsal view, apical margins 
rounded, inner apical angle not prominent, situated pos- 
terior level of outer angles, sutural margin not raised, 
sutural striae deep, slightly converging apically, not 
curved at base and not extending laterad pronotal lobe, 
adsutural areas flat. Elytral punctation very fine and 
sparse on basal third, becoming more distinct in mid- 
dle third and consisting of poorly defined, very shallow, 
dense and fine punctures, inconspicuous on apical third 
of elytra. Hypomera smooth. Mesepimera about 4 times 
as long as wide and 1.5 times as long as interval to 
mesocoxa. Metaventrite slightly flattened basomedially, 
lacking grooves or impressions, without microsculpture, 
very finely punctate. Submesocoxal areas 0.04 mm, 
about as third of interval to metacoxae, submesocoxal 
lines convex. Metanepisternum flat, hardly narrowed 
anteriad, with inner margin weakly rounded, somewhat 
impressed below margin of metaventrite. Tibiae straight. 
Abdomen with microsculpture consisting of transverse 
striae. Sternite 1 very finely punctate. Submetacoxal 
areas 0.06-0.07 mm, almost as half of interval to apical 
margin of sternite, submetacoxal lines convex, with fine 
margin punctures. 

Male: Protarsal and mesotarsal segments | to 3 slightly 
widened. Aedeagus (Figs 99-102) 0.78-0.83 mm long, 
fairly strongly sclerotized, symmetrical. Median lobe 
with basal bulb large, longer than apical process, articu- 
lar process inconspicuous. Ventral branch of apical pro- 
cess inflexed, with ventral side sinuate, subapical dorsal 
tooth followed by very fine dorsal crenulations, tip in 
dorsal and lateral views acute. Dorsal branches of api- 
cal process very short. Parameres fairly narrow in basal 
halves, each with large weekly sclerotized subapical lobe 
distinct in dorsal view, abruptly widened in apical halves 
in lateral view. Internal sac complex, with basal tuft of 
very dense spine-like structures, two admesal groups of 
large, curved denticles, followed by irregular rows of 
fairly robust mesal denticles. Membranes with scale-like 
structures. 


Distribution: Indonesia: Sumatra, Lombok. 


110 I. Lobl 


Fe 
t 
ENT 
A 
Li 


IN 


ZI 


Figs 95-100 

(95-98) Scaphisoma aspectum sp. nov., median lobe of aedeagus and parameres, without proximal part of basal bulb (95) in dorsal 
view, proximal half of internal sac (96) and entire internal sac (97) in dorsal view, median lobe of aedeagus and parameres, 
without proximal part of basal bulb (98) in lateral view view; scale bars = 0.2 mm. 

(99-100) Scaphisoma gracilicorne Achard, aedeagus (99) in dorsal view and internal sac in dorsal view (100). 

Scale bars = 0.2 mm (95, 97, 99) and = 0.1 mm (96, 100). 


Scaphidiinae of the Lesser Sunda Islands 111 


Comments: The species is a member of the Scaphisoma 
haemorrhoidale group, well characterized by its aedea- 
gus. Achard (1920) failed to give any statement about 
the number of specimen examined and their deposi- 
tory. The specimen designated here as lectotype bears 
his original identification label and complies with his 
description except for the length he gave as | mm. 


Scaphisoma jacobsoni Löbl, 1975 


Additional material examined: SMNS, MHNG; 
31 ex.; Indonesia, Bali 12 km NW of Bedugul (Buyan 
lake) 29.Apr.-2.May 2001 Bolm Igt., 900 m. 


Distribution: Indonesia: Sumatra, Java, Bali; East 
Malaysia: Sarawak; Thailand. 


Comments: The species may be easily distinguished by 
its elytra with apically strongly converging sutural striae 
and very fine discal punctation, in combination. The api- 
cal process of the median lobe is trilobed and symmetri- 
cal, as in members of the S. haemorrhoidale group, but 
the dorsal lobes are weakly sclerotized and concealed. 


Scaphisoma luteomaculatum Achard, 1920 
Figs 103-106 


Scaphosoma luteomaculatum Pic, 1915b: 5. 

Scaphosoma sapitense Pic var. infasciatum Achard, 1920: 131, 
syn. nov. 

Scaphisoma dansalanense Löbl, 1972: 95, syn. nov. 


D 


Lectotype of /uteomaculatum Pic: MNHN; 6; by 
present designation, labelled: Java merid. Palabuan 
1892 H. Fruhstorfer. (printed) / type (handwritten by 
Pic) /TYPE (red, printed) / n. sp. (handwritten by Pic) 
/ Scaphosoma (handwritten by Pic) / Scaphosoma 
luteomaculatu Pic (handwritten by Pic) / Lectotype 
Scaphisoma luteomaculatum Pic det. Löbl, 2014. The 
aedeagus is mounted on a celluloid slide. 


Lectotype of infasciatum Achard: MNHN; ©; by 
present designation, labelled: Lombok Sapit 2000’ 
April 1896 H. Fruhstorfer /Museum Paris Coll. 
A. Grouvelle 1915 /S. sapitense Pic v. infasciatum 
TYPE (handwritten by Achard) J Achard det / TYPE 
(red) / Lectotype Scaphisoma infasciatum Achard det. 


Löbl, 2014. 


Additional material examined: NMPC, MHNG; 9 
ex.; with the same data as the lectotype. - SMNS; 1 9; 
Lombok, Sesaot, 500 m, 1.2.1994 Bolm Igt. - NMPC; 
1 3; Lombok, Pringabaja, April 1896 H. Fruhstorfer. 
— SMNS, MHNG; 3 €, 3 ©; Lombok, Senaro N slope 
of Rinjani, 2.-5.Feb.1994 1100 m Bolm Igt. —- SMNS; 
1 ©; E Lombok, Sapit, 14.-16.2.1994 SE slope of Mt. 
Rinjani Bolm Igt. - MHNG; 1 ©; Lombok, Pusut Pass 
300 m, 3.XI.1991, I. Löbl. - SMNS, MHNG; 7 <, 
5 2;W Sumbawa, Batudulang, 30 km S of Sumb. Besar, 


1000 m Bolm legt. 10.Feb 1994. — SMNS, MHNG; 
1 &, 4 ©; NE Sumbawa, Calabai (Tambora N.P.) 11.- 
13.Feb.1994, Bolm Igt. - MHNG; 2 d; Bali, Lake 
Dratan, 24.1V.81, Rougemont. -— MHNG; 2 <; Bali, 
Gunung Agung, 19.VII.82, de Rougemont. -— MHNG; 
1 ©; Sitinjaulaut, alt. 1000 m 25 km east from Padang 
Sumatera Barat VIII.17.1977, Sinji Nagai leg. 


Distribution: Philippines: Mindanao; Indonesia: 
Sumatra, Java, Bali, Lombok, Sumbawa, Buru. 


Comments: The colour pattern of the elytra is quite 
variable. Extreme are specimens with elytra uniformly 
light reddish, or specimens having elytra blackish with 
a well delimited large reddish subhumeral spot and 
light in apical third. The aedeagus (Figs 103-106) has a 
complex internal sac, and the shape of the parameres is 
slightly variable. The record from Burma (Myanmar) by 
Pic (1921) was not yet verified. 


Scaphisoma lombokense Löbl, 1986 
Distribution: Indonesia: Lombok. 


Comments: The description of this species was 
based on a single specimen from Sesaot (in MHNG). 
Additional material is unknown. 


Scaphisoma obliquemaculatum Motschulsky, 1863 


N 


Additional material examined: SMNS, MHNG; 2 4, 
1 ©; Indonesia, W. Sumbawa Batudulang, 30 km S of 
Sumb. Besar, 1000 m Bolm Igt. 10. Febr. 1994. 


Distribution: Tropical Asia from Sri Lanka to Vietnam, 
Great Sunda Islands, Lesser Sunda Islands: Sumbawa, 
Mascarene Archipelago. 


Comments: The species was redescribed and its 
aedeagus illustrated in Löbl, 1971. The female from 
Batudulang has a reddish anterior part of the elytron 
clearly separated from yellowish elytral apex by a dark 
transverse band, as other specimens coming from diffe- 
rent areas. The two males from Batudulang differ by the 
elytra uniformly dark from their bases upto the yellow- 
ish apical bands. 


Scaphisoma rufescens (Pic, 1920) 


Additional material examined: SMNS; 1 4; 
Indonesia, Bali, Danau Buyan, 1300 m, 19.-21.2.1994, 
Bolm Igt. 


Distribution: South-east Asia, from Yunnan, Vietnam, 
Thailand, Malaysia, Singapore to Indonesia: Kalimantan 
and Bali. 


Comments: The species was redescribed and its aedea- 
gus illustrated in Löbl, 1981b. 


11122 I. Löbl 


js 
N 
SION 


n (C 
PI 7 
447, : a N 
‘493, SAN NER 
DOS D Parents 
AAT Ds AN d STR vb 
DI NA (MIA Dare N 
SINS DERMI DMN 
7749 NOS) Vd a ah NS 
ZUG NF ANAC A i 
ZA4, Sy Vln me a SUN 
7 71,9 CS BEA Y PRAR 
OR OR 06 103 
77 > N SS dI ‘1 

4) )) 2 RES ‘ ‘ 

ADS SEE 


104 105 106 107 


Figs 101-107 

(101-102) Scaphisoma gracilicorne Achard, aedeagus, without proximal part of basal bulb (101) in lateral view, paramere in ventral 
view (102). 

(103-106) Scaphisoma luteomaculatum Pic, internal sac (103) in dorsal view, (104-106) parameres in ventral view, from Bali (104), 
Buri (105) and Mindanao (106); scale bars = 0.1 mm. 

(107) Scaphisoma sapitense Pic, median lobe and internal sac in dorsal sac, without proximal part of basal bulb; scale bar = 
0.2 mm. 


Scaphidiinae of the Lesser Sunda Islands 113 


Scaphisoma sapitense Pic, 1915 
Figs 107-111 


Scaphosoma sapitense Pic, 1915a: 15. 


Lectotype: MNHN; ©; by present designation, labelled: 
Lombok Sapit 2000’ April 1896 H. Fruhstorfer/TYPE 
(red)/ Scaphosoma sapitense Pic (handwritten by Pic)/ 
Lectotype Scaphisoma sapitense Pic det. Löbl, 2014. 


Additional material examined: NMPC, MHNG; 
10 ex.; with the same data as the lectotype. — SMNS, 
MHNG; 27 ex.; Lombok, Sapit-Sembalun Bumbung 
14.-16. Feb 1994 900-1500 m, Bolm Igt. — SMNS, 
MHNG; 2 ex.; Lombok, Sapit SE slope of Mt. Rinjani, 
14.-16.Feb 1994 Bolm Igt. - SMNS, MHNG; 17 ex.; 
Lombok, Senaro, N slope of Rinjani, 2.-5. Feb. 1994 
1100 m Bolm Igt. 


Redescription: Length 1.90-2.15 mm, width 1.25- 
1.35 mm. Head, pronotum and venter of thorax dark 
reddish-brown to blackish. Elytra light reddish-brown 
to yellowish, with darkened bases and adsutural areas, 
each elytron with dark brown to black discal spot. 
Latter variable in size, usually not touching sutural stria. 
Abdomen with basal sternites usually dark brown, api- 
cal sternites, exposed tergites and appendages about 
as light as prevailing elytral surface. Pronotum and 
elytra lacking microsculpture. Antennae long, length/ 
width ratio of antennomeres as follows: III 14/9: IV 
36/7: V 50/8: VI 55/8: VII 52/14: VII 45/11: IX 55/14: 
X 52/15: XI 63/15. Antennomere III subtriangular. 
Pronotum finely punctate, with lateral margins evenly 
rounded, lateral margin carinae visible near base or in 
basal half in dorsal view. Apex of scutellum exposed. 
Elytra with lateral margin carinae entirely exposed in 
dorsal view, apical margins truncate, inner apical angle 
rectangular, not prominent in male, prominent and 
tooth-like in female, situated posterior level of outer 
angles, sutural margin not raised, sutural striae deep, 
parallel, at base somewhat curved, not extending laterad 
pronotal lobe, adsutural areas flat. Elytral punctation 
fairly coarse, dense, punctures well delimited, puncture 
intervals mostly about as large to twice as large as punc- 
ture diameters. Hypomera smooth. Mesepimera longer 
than interval to mesocoxa. Metaventrite microsculptured 
on areas between submesocoxal lines and metacoxae, 
with median area slightly convex, two indistinct apico- 
median impression, punctation on apicomedian area fine 
but distinct, on lateral areas very fine. Submesocoxal 
areas 0.04 mm, about as fifth of intervals to metacoxae, 
submesocoxal lines subparallel, with coarse margin 
punctures. Metanepisternum flat, hardly narrowed ante- 
riad, with inner margin almost straight, impressed below 
margin of metaventrite. Tibiae straight. Abdomen with 
microsculpture consisting of transverse striae. Sternite 
1 with distinct mediobasal punctation, very finely 
punctate on remaining surface. Submetacoxal areas 
0.06-0.07 mm, submetacoxal lines convex, with coarse 
margin punctures. 


Male: Protarsal and mesotarsal segments | to 3 distinctly 
widened. Inner apical angle of elytra not prominent. 
Aedeagus (Figs 107-111) 1.30-1.37 mm long. Median 
lobe and parameres symmetrical, strongly sclerotized. 
Median lobe with large basal bulb, ventral branch of api- 
cal process short, in apical half weakly inflexed, with sub- 
apical dorsal denticle, dorsal branches of apical process 
slightly shorter than ventral branch, articular process not 
prominent. Parameres in axis with median lobe, slightly 
sinuate and narrowed apically in lateral view, gradually 
narrowed toward level of tip of median lobe in dorsal 
view and with finely crenulated inner margin; pores scat- 
tered, situated in basal halves. Internal sac complex, with 
tuft of dense denticles, overlapping median row of scle- 
rotized scales, membranes with scale-like structure. 
Female: Inner apical angle of elytra prominent, tooth- 
like. 


Distribution: Indonesia: Lombok. 


Comments: The original description, based on speci- 
men/s from Lombok: Sapit, gives only the colour pat- 
tern and body length. Achard (1920: 131) reported the 
species from “Lombok: Sapit, altitude 2000 mètres et 
Pringabaja (H. Frühstorfer, avril 1896)”. In fact, the 
altitude on the labels is in feet, and the specimen from 
Pringabaja was misidentified. Achard (1. c.) mentioned 
also the shape of the sutural striae and of the 3rd anten- 
nomere, found the colour pattern variable, and described 
a new variety, infasciatum, distinguished from the 
nominal form by the elytra lacking a dark discal spot. 
His infasciatum is a distinct species and his statement 
about the variable colour in S. sapitense is erroneous. 
Specimens with the same locality labels as the lecto- 
type and housed in the NMPC are possibly syntypes but 
not considered as such, in absence of evidence that they 
were used by Pic for the description. 


Scaphisoma sesaotense Löbl, 1986 


Additional material examined: SMNS, MHNG; 2 4, 
1 ©; Indonesia, NE Sumbawa, Calabai (Tambora N.P.) 
11.-13.Febr.1994 Bolm Igt. 


Distribution: Indonesia: Lombok, Sumbawa. 


Comments: One of the males and the female from 
Calabei have elytra darkened along their bases, the 
sutural striae and on a narrow transverse band, while the 
other male has the elytra uniformly reddish-brown, as 
mentioned in the description of this species, based on a 
single specimens from Lombok, Sesaot. 


Scaphisoma sp. 


Material examined: SMNS; 1 ©: Indonesia, Bali 
12 km NW of Bedugul (Buyan lake) 29.Apr.-2.May 
2001 Bolm lgt., 900 m. 


114 I. Lobl 


€ 


nia 


SS 


= TER IT 
D N SKK 
N o RT = 
SNS 
AEOSZ 
= 


SS 


3 SA 
DA SN 
A SSS 
ry — > > 
SIAE 
È = SS 
wy Sec 
3 = 
SEE 


115 


114 


113 


Figs 108-115 
(108-111) Scaphisoma sapitense Pic, aedeagus (108) in dorsal view; scale bar = 0.2 mm, apical half of internal sac (109) in dorsal 
view; scale bar = 0.1 mm, median lobe and parameres (110), without proximal part of basal bulb, in lateral view, paramere 


(111) in dorsal view, without its base; scale bars = 0.2 mm. 
(112-113) Scaphobaeocera baliensis sp. nov., aedeagus in dorsal (112) and ventral (113) views. 
(114-115) Scaphobaeocera lombokensi sp. nov., aedeagus in dorsal (114) and ventral (115) views; scale bars= 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 115 


Comments: This is one of the larger Indonesian species 
of Scaphisoma. It is 1.90 mm long, 1.25 mm wide and 
has conspicuous colour pattern and antennae: the elytra 
are as the pronotum very dark around margins, most of 
the elytral disc is covered by light, reddish, somewhat 
oblique spot. The antennomeres III and IV are minute, 
much narrower, and combined much shorter than V. 
In addition, the species is characterized by the sutural 
striae of elytra extending along bases to form basal 
striae reaching beyond elytral basal width, in combina- 
tion with very finely punctate ventral side of the body 
and strigulate abdominal microsculpture. 


Scaphobaeocera Csiki, 1909 


Comments: Currently, 94 species of Scaphobaeocera 
are recognized as valid, most of them occurring in 
tropical and subtropical Asia. The range of the genus 
is, however, extended northwardly to Far East Russia, 
and to Queensland and Subsaharian Africa in the south. 
Scaphobaeocera kraepelini (Pic, 1933) from Java is the 
sole so far mentioned Indonesian species, and for prac- 
tical reasons redescribed below. Most members of this 
genus possess elytra with distinct parasutural striae, 
unknown in other scaphidiines, any many have elytra, 
eventually also other parts of the body, iridescent, a 
character state rather unusual in other genera of the sub- 
family. 


Scaphobaeocera baliensis sp. nov. 
Figs 112-113 


Holotype: MHNG;  ; Indonesia, Bali, Mt. Batukaru nr. 
Luhur Temple, 500-700 m, I. Löbl, 28-29.X.1991. 


Etymology: The species epithet is derived from the 
name of the island Bali. 


Description: Length 1.10 mm, width 0.56 mm, dor- 
soventral diameter 0.67 mm. Head and most of body 


very dark reddish-brown, thorax almost blackish. 
Abdomen somewhat lighter than thorax and elytra. 
Femora and tibiae reddish-brown, tarsi and antennae 
lighter. Thorax and elytra lacking obvious microsculp- 
ture, hardly iridescent. Pronotal and elytral punctation 
very fine, elytral punctures slightly coarser than pro- 
notal. Length/width ratio of antennomeres as follows: 
III 16/6: IV 19/6: V 26/6: VI 18/6: VII 35/9: VIII 15/8: 
IX 40/11: X 40/12: XI 50/13. Minute tip of scutellum 
exposed. Elytra with sutural striae starting at margin of 
pronotal lobe, parasutural striae very fine but distinct. 
Hypomera without striae. Middle part of metaventrite 
convex, lacking impression or groove, smooth on nar- 
row anteriomesal area, densely and distinctly punc- 
tate and with short pubescence on lateromesal areas 
and entire posterior half. Lateral parts of metaven- 
trite lacking microsculpture, very finely punctate, with 
sparse, short pubescence. Submesocoxal areas about 
0.02 mm long, submesocoxal lines with very fine mar- 
ginal punctures not extending along mesepimera. 
Metanepisternum flat, parallel-sided, 0.06 mm wide. 
Tibiae straight. Abdominal sternite 1 with fairly coarse, 
elongate basal punctures, remaining punctation very 
fine. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
112-113) 0.36 mm long. Median lobe narrow, with short, 
hardly inflexed apical process. Ventral side of apical pro- 
cess weakly concave. Articular process of median lobe 
indistinct. Parameres very narrow in basal two thirds, dis- 
tinctly widened in apical third (lateral view), widest sub- 
apically, about in level with tip of median lobe; straight 
but at apices curved mesally in dorsal view. Internal sac 
with flagellum fairly wide, forming four complete loops, 
slightly narrowed posterior basal loop. 


Distribution: Indonesia: Bali. 


Type locality: Bali, Mt. Batukaru nr. Luhur Temple, 
500-700 m. 


Comments: The aedeagal characters suggest relation- 
ships between S. baliensis and S. kraepelini from Java, 


Key to the Scaphobaeocera species of the Lesser Sunda Islands 


I 


Antennomere VII more than twice as long as VIII, antennomere XI almost 4 times as long as wide. Aedeagus with 
Hasellumgspiralgtor min PHOUNRCOMPIELEILOO DS nn S. baliensis sp. nov. 
Antennomere VII less than twice as long as VIII, antennomere XI about 2 to 3 times as long as wide. Aedeagus 
with flagellum spiral and forming single complete loop, or not spiral................................. 2 
Apicomedian part of metaventrite with sparse pubescence, middle ofmetaventrite smooth. Aedeagus with flagellum 
SHOTLSTIOUISPIEA Eee ER ee species near S. werneri Löbl 
Entire median part of metaventrite with dense and short pubescence. Aedeagus with long flagellum.................... 3 
Antennomere VIII twice as long as wide. Median lobe of aedeagus with large, prominent articular process........... 
ee NEN nS ee ee LATI IIIZII e ee tiens traces sauts ses nte nee se saute 05 S. pseudotenella Löbl 
Antennomere VIII about 2.5 times as long as wide. Median lobe of aedeagus without prominent articular process 


Aedeagus with flagellum filiform, forming loop posterior mid-length ...................... S. bulbosa Löbl 
Aedeagus with flagellum proximally widened, forming basal 100p.................................... S. lombokensis sp. nov. 


116 


S. confusa Löbl, 1986 from Queensland, and S. spira 
Löbl, 1990 from Thailand. These three species share 
comparatively short apical process of the median lobe, 
apically widened parameres (in lateral view) and spiral 
flagellum forming multiple loops. The new species may 
be distinguished by the thicker flagellum forming only 
four complete loops, the apical process of the median 
lobe being almost in axis with the basal bulb, and the 
parameres with the widest points situated about inlevel 
with the tip of the median lobe. In addition, S. baliensis 
differs from its allied by the comparatively longer api- 
cal antennomeres. Scaphobaeocera dorsalis Löbl, 1980, 
widely distributed in Southeast Asia, possesses also 
a flagellum forming four loops. It differs by the much 
broader flagellum, the apical process of the median lobe 
inflexed, and in external characters by the hypomera 
each with a longitudinal stria. 


Scaphobaeocera bulbosa Löbl, 2011 


Material examined: MHNG; 18 ©, 20 ©; Indonesia, 
Bali, Mt. Batukaru nr. Luhur Temple, 500-700 m 
I. Löbl, 28-29.X.1991. 


Distribution: Philippines: Luzon; Indonesia: Bali. 


Comments: All males were dissected and their aedeagi 
examined. Several additional males in poor condition 
remain unidentified. 


Scaphobaeocera lombokensis sp. nov. 
Figs 114-115 


Holotype: MHNG; <; Indonesia, Lombok Pusuk Pass, 
300 m 3.XI.1991, I. Löbl degr. forest leaf litter. 


Paratypes: MHNG; 1 £, 3 ©; with the same data as the 
holotype. 


Etymology: The species epithet is derived from the 
name of the island Lombok. 


Description: Length 1.05-1.12 mm, width 0.53-0.55 
mm, dorsoventral diameter 0.62-0.64 mm. Head and 
body fairly light reddish-brown, mesoventrite and 
metaventrite eventually darkened. Apex of abdomen, 
femora and tibiae slightly lighter, tarsi and anten- 
nae much lighter. Thorax and elytra lacking obvious 
microsculpture, not iridescent. Pronotal and elytral 
punctation similar, very fine. Length/width ratio of 
antennomeres as follows: III 14/5: IV 19/5: V 22/5: VI 
20/5: VII 21/7: VIII 15/6: IX 24/9: X 24/10: XI 35/11. 
Scutellum concealed. Elytra with sutural striae starting 
at margin of pronotal lobe, parasutural striae indistinct. 
Hypomera without striae. Middle part of metaventrite 
flat, lacking impression or groove, entirely densely and 
distinctly punctate and with short pubescence. Lateral 
parts of metaventrite lacking microsculpture, very finely 


I. Löbl 


punctate, with sparse, short pubescence. Submesocoxal 
areas about 0.03 mm long, submesocoxal lines impunc- 
tate. Metanepisternum flat, parallel-sided, 0.03 mm 
wide. Tibiae straight. Abdominal sternite 1 without dis- 
tinct basal punctures, punctation very fine. 

Male: Protarsomeres | to 3 distinctly widened. Aedeagus 
(Figs 114-115) 0.27 mm long. Median lobe narrow, with 
long, moderately inflexed apical process. Apical process 
tapering in lateral view, with ventral side weakly con- 
cave. Articular process of median lobe small, not promi- 
nent. Parameres narrow between bases and apical halves, 
widened in apical halves and weakly sinuate in lateral 
view, with widest points being about in level with tip of 
median lobe; almost straight in dorsal view. Internal sac 
with flagellum fairly wide in proximal part, forming sin- 
gle complete loop, gradually narrowed apically. 


Distribution: Indonesia: Lombok. 
Type locality: Lombok Pusuk Pass, 300 m 


Comments: The aedeagal characters suggest rela- 
tionships with S. curvipes Löbl, 1977 from Australia, 
S. fratercula Löbl, 1984, S. molesta Lôbl, 1999 from 
China, S. variabilis Löbl, 1981 from Japan and S. mus- 
sardi Löbl, 1971 known from the Indian subcontinent. 
Scaphobaeocera curvipes and S. mussardi possess fla- 
gellum forming two loops, and the parameres in the for- 
mer species bear each a ventral membranous lobe, while 
they are gradually widened apically in the latter species. 
The new species may be readily distinguishes from 
S. curvipes by its smaller body-size and straight tibiae. 
Scaphobaeocera variabilis is distinctly microsculptured 
and iridescent dorsally and ventrally, has parameres 
widened in apical two thirds and flagellum forming 
three loops. Scaphobaeocera fratercula has the flagel- 
lum forming a single, incomplete loop and differs nota- 
bly by its elytra with distinct parasutural striae and by 
the impunctate and grooved centre of the metaventrite. 
The flagellum in Scaphobaeocera molesta is marrow 
with abruptly widened base; in addition this species has 
a large and iridescent body, striate hypomera, and elytra 
with distinct parasutural striae. 


Scaphobaeocera pseudotenella Löbl, 2011 


Material examined: MHNG; 2 ; Indonesia, Bali, 
Mt. Batukaru nr. Luhur Temple, 500-700 m I. Löbl, 
28-29.X.1991. 


Distribution: Philippines: Luzon; Indonesia: Bali. 


Comments: The parameres are broader in the two spe- 
cimens from Bali than in those from Luzon (see Löbl, 
2011: 712). More material would be needed to assess 
the variability of the character. 


Scaphidiinae of the Lesser Sunda Islands 117 


Scaphobaeocera sp. 


Material examined: MHNG; 1 4, 1 9; Indonesia, Bali 
Lake Buyan ca 1200 m 8.-9.X1.1991, I. Löbl degr. forest 
floor litter. 


Comments: The single available male has its aedeagus 
deformed, with the internal sac extruded. It possesses 
characters of S. werneri Löbl, 2011 from Mindanao but 
the narrow apical part of the flagellum is shorter and the 
metanepisterna are broader. 


Scaphobaeocera kraepelini (Pic, 1933) 
Figs 116-117 


Toxidium kraepelini Pic, 1933: 72. 


Lectotype: MNHN; d; by present designation, labelled: 
Buitenzorg, Java K. Kraepelin leg. 24.11.-12.111.1904 
ded. 8. VI.1904 (printed) / T. Kraepelini desire (hand- 
written by Pic) / Scaphobaeocera kraepelini (Pic) 
det. Lôbl 1977 / Lectotype (red, printed) / Lectotype 
Toxidium kraepelini Pic det. Löbl, 2014. 


Redescription: Length 1.05 mm, width 0.62 mm, 
dorsoventral diameter 0.64 mm. Head and body light 
reddish-brown, mesoventrite and metaventrite slightly 
darkened. Abdomen, femora and tibiae slightly lighter, 
tarsi and antennae yellowish. Thorax and elytra lacking 
obvious microsculpture, not iridescent. Pronotal and 
elytral punctation similar, very fine, hardly visible at 
100x magnification. Length/width ratio of antennomeres 
as follows: VIII 20/6: IX 28/11: X 30/12: XI 42/14 
(antennomeres III to VII missing). Minute point of 
scutellum exposed. Elytra with sutural striae starting 
at margin of pronotal lobe, parasutural striae absent. 
Hypomera lacking striae. Middle part of metaventrite 
convex, lacking impression or groove, entirely densely 
and distinctly punctate, bearing short pubescence. 
Lateral parts of metaventrite lacking microsculpture, 
very finely punctate, with sparse, short pubescence. 
Submesocoxal areas hardly 0.02 mm long, about as 
fourth of interval to metacoxae. Submesocoxal lines 
parallel, with coarse, elongate marginal punctures. 
Metanepisternum flat, large, about 0.07 mm wide, 
slightly narrowed apically, strongly narrowed anteriad, 
with deep, convex suture. Tibiae straight. Abdominal 
sternite 1 with striagulate microsculpture, and coarse, 
elongate basal punctures, remaining punctation very 
fine. 

Male: Protarsomeres hardly widened. Aedeagus (Figs 
116-117) 0.25 mm long. Median lobe gradually nar- 
rowed, with apical process short, hardly inflexed, very 
narrow at tip in lateral view. Articular process of median 
lobe indistinct. Parameres irregularly bent and strongly 
narrowed toward apices. Internal sac with narrow flagel- 
lum forming four complete loops. 


Distribution: Indonesia: Java. 


Comments: M. Pic usually failed to mention numbers 
of specimens he examined, nor to designate holotypes. 
This is also true for Toxidium kraepelini as for the other 
taxa described in the same paper (Pic, 1933). However, 
Pic stated that “cotypes” of three species, including of 
Toxidium kraepelini, are in his collection, actually in 
MNHN, while other specimens belong to the collec- 
tion of the Zoological Museum, Hamburg. According to 
Horn et al. (1990), the latter collection was destroyed in 
1943. Thus, the specimen of Toxidium kraepelini housed 
in MNHN is the sole syntype actually preserved and is 
here designated as lectotype. Its characters and label 
data fit the Pic’s description. . 

The species was transferred to Scaphobaeocera in Löbl, 
1984: 82, but not redescribed. 


Scaphoxium Löbl, 1979 


Comments: The range of this genus is similar to that 
of Scaphobaeocera, though it is with 41 species cur- 
rently recognized less species-rich and usually poorly 
represented in collections. The genus was not yet 
reported from Indonesia. Scaphoxium may be readily 
distinguished from other scaphidiines having strongly 
approximate middle and hind coxae by the curved 3rd 
antennomere and lobed ventral margins of the hypo- 
mera. 


Scaphoxium bilobum sp. nov. 
Figs 118-120 


Holotype: MHNG; <; Indonesia, Lombok, Mt. Rinjani, 
ca 400 m nr. Waterfalls, 5.XI.91 Löbl, veg. debris nr. 
river. 


Paratypes: MHNG; 3 &; Indonesia, Lombok Mt. 
Rinjani above Senaro 900-1100 m 6.X1.1991, I. Löbl 
forest floor litter. - MHNG; 1 ©; Lombok Batu Coq 
(N of G. Rinjani) 500 m, sec. forest in gorge D. Agosti, 
12.03.1991. 


Etymology: The species epithet is a Latin adjective 
referring to the lobed parameres. 


Description: Length 1.38-1.45 mm, width 0.65- 
0.67 mm, dorsoventral diameter 0.74-0.76 mm. Body 
fairly dark reddish-brown. Femora and tibiae slightly 
lighter, apex of abdomen, tarsi and antennae light, 
almost yellowish. Length/width ratio of antennomeres 
as follows: III 17/5: IV 16/4: V 25/5: VI 35/6: VII 35/9: 
VII 28/10: IX 33/10: X 30/10: XI 38/10. Pronotal 
punctation very fine, hardly visible at 50x magnifica- 
tion. Scutellum concealed. Elytral punctation some- 
what less fine than pronotal punctation; sutural striae 
starting about 0.15 to 0.25 mm behind margin of pro- 
notal lobe. Mesoventrite with narrow median groove, 
not striate. Mesepisterna very finely punctate. Median 


118 I. Löbl 


124 


120 121 122 123 


Figs 116-124 

(116, 117) Scaphobaeocera kraepelini (Pic), aedeagus in dorsal (116) and ventral (117) views. 

(118-120) Scaphoxium bilobum sp. nov., aedeagus (118) in dorsal view, internal sac (119) in dorsal view, paramere (120) in ventral 
view. 

(121-124) Xotidium bolmarum sp. nov., aedeagus in dorsal (121) and ventral (122) views, (123) median lobe, without basal bulb, and 
paramere in lateral view, (124) gonocoxite. 

Scale bars = 0.1 mm. 


Scaphidiinae of the Lesser Sunda Islands 119 


part of metaventrite slightly convex, flattened near 
metacoxal process, very finely punctate. Lateral parts 
of metaventrite with several comparatively coarse punc- 
tures, impunctate near apical margin. Submesocoxal 
areas about 0.05 mm long, shorter than interval to 
metacoxae. Submesocoxal lines coarsely punctate. 
Metanepisternum flat, 0.05 mm wide, with straight 
suture. Abdomen with distinct punctate microsculpture, 
basal punctures on sternite | distinct, remaining abdom- 
inal punctation hardly visible. 

Male: Protarsi hardly widened. Aedeagus (Figs 118-120) 
0.41-0.44 mm long. Median lobe with acute tip. Para- 
meres narrowed between bases and subapical lobe, latter 
rounded, with strongly narrowed apical section about 
as long as fourth of parameral length. Internal sac with 
pair of mesal rods and pair of admesal plates tapering 
proximally, membranes lacking denticulate or spine-like 
structures. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok, Mt. Rinjani, ca 400 m nr. 
Waterfalls. 


Comments: Following congeners possess lobed par- 
ameres and internal sac bearing two pairs of sclerites: 
S. oxyurum (Löbl, 1977) and S. cuspidatum (Löbl, 1977) 
from Australia, S. keralense Löbl, 1979 from India, 
S. taiwanum Löbl, 1980 from Taiwan, and S. avidum 
Lôbl, 1990 and S. topali Löbl, 1981 from Vietnam. 
The latter species differs drastically by the parameres 
abruptly widened posterior subapical lobes. Scaphoxium 
cuspidatum, S. oxyurum, and S. avidum may be distin- 
guished from S. bilobum by the internal sac having an 
additional small sclerite. The sclerotized pieces of the 
internal sac are very small and distant in S. keralense, 
and this species has also denticulate membranes in the 
middle part of the internal sac. Finally, S. bilobum may 
be easily distinguished from S. taiwanum by the nar- 
rower parameres and the proximally strongly narrowed 
admesal pair of sclerites of the internal sac. 


Key to the species of Xotidium 


Xotidium Lôbl, 1992 


Comments: This genus is characterized by 2-segmented 
labial palpi. Currently, it includes six species, known 
from Sri Lanka, Nepal Himalaya, the Philippines, 
Australia, and Mauritius. An additional species was 
found in Lombok. 


Xotidium bolmarum sp. nov. 
Figs 121-124 


Holotype: SMNS; &; Indonesia, Lombok Senaro, N 
slope of Rinjani, 2.-5.Feb.1994 Bolm Igt. 1100 m. 


Paratypes: SMNS, MHNG; 3 7, 8 2; Indonesia, with 
the same locality data as the holotype. - SMNS; 1 4; 
Lombok Is. Sapit-Sembalun Bumbung 14.-16.Feb.1994 
Bolm Igt. 900-1500 m. 


Etymology: The species epithet is patronymic. 


Description: Length 1.35-1.46 mm, width 0.80- 
0.85 mm, dorsoventral diameter 0.78-0.80 mm. Body 
reddish-brown, apex of abdomen yellowish, femora and 
tibiae somewhat lighter than pronotum and elytra, tarsi 
and antennae yellowish-brown to yellowish. Antennae 
long, length/width ratio of antennomeres as follows: III 
29/6: IV 33/6: V 38/7: VI 40/6: VII 48/12: VII 45/7: 
IX 46/10: X 43/13: XI 44/15. Dorsal surface of body 
very finely punctate. Elytra with sutural striae shallow, 
curved along bases to form basal striae joined to lateral 
striae. Adsutural areas flat. Mesal part of metaventrite 
flattened. Submesocoxal areas about 0.05 mm long, 
submesocoxal lines coarsely punctate. Remainder of 
metaventrite very finely punctate. Abdomen without 
obvious microsculpture, with fairly coarse basal punc- 
tures, remaining punctation hardly visible. 

Male: Male protarsus hardly widened. Aedeagus (Figs 
121-123) 0.45-0.48 mm long. Widened apical part of 
parameres short, bent dorsally, with weakly sclerotized 


l Blytrallackinp4basalistuaeSsuturalistriaefShOKeNed eee O X. mauritianum (Vinson) 
= BW Wallin LAsgll sims jo med fo suina S111EYS.cc000000000c00000000s09000056e05000000000000000000000000000000080000000000000000300000005000803200000000 2 
2 Bilytragunitormlygreddish-DLow 1 O1 AC RER O into 3 
= BR Vin GSMO. COIOUT TELE .003000090000000000600000000040 I DOME COL PURE EEE EEE PE CEE Eee Ee 6 
3 Body length 1.10-1.25 mm. Aedeagus with short, evenly thick and sinuate flagellum ............ X. pygmaeum (Löbl) 
— Bodyalen theless Salt spin cognac nan DRE RER D EE PE EEE E ES ERP 4 

Widened apical section of parameres about as long as half of parameres. Internal sack with very long, weekly 

Sclerotizedtandisim pleitu DERE A RE O O I ir X. tubuliferum Löbl 
= Widened apical section of parameres much shorter than half of parameres .…....................................................... 5 
5 Internal sac of aedeagus with U-shaped, basally thickened flagellum .............. X. uniforme Lobl 
= Internal sac of aedeagus with short basal sclerites followed by long membranous tube ......... X. bolmarum sp. nov. 
6 Elytra dark, each with light transverse fascia situated in basal half of elytron and light apical part. Pronotum 


entirelyidarkesnsege ron nananana 


D SR oO ci X. montanum (Löbl) 


= Elytra light, each darkened along basal and apical margins, usually also darkened along sutural margin. Pronotum 


light, usually with dark transverse fascia.................... 


WR IT X. notatum (Löbl) 


margins. Internal sac with short basal sclerites followed 
by membranous tube bearing very fine spine-like struc- 
tures. 

Female: Ovipositor (Fig. 124) with long distal gonoco- 
xite slightly narrowed between base and mid-length, 
bearing one short subapical seta, and one long and one 
short apical setae, gonostyle absent. 


Distribution: Indonesia: Lombok. 


Type locality: Lombok Senaro, N slope of Rinjani, 
1100 m. 


Comments: This new species may be easily distin- 
guished from other congeners by characters given in 
the key. It is unambiguously defined by the shape of the 
parameres in combination with the sclerotized structures 
of the internal sac. 


ACKNOWLEDGEMENTS 


Following colleagues kindly provided material examined 
in the present study: D. Agosti, Bern, T. Deuve, Paris, J. P. 
Duffels, Amsterdam, J. Häjek, Praha, G. de Rougemont, 
London, W. Schawaller, Stuttgart, and A. Taghavian, 
Paris. In addition, the assistance of A. Cibois, Genève, is 
gratefully acknowledged. 


REFERENCES 


Achard J. 1920. Notes sur les Scaphidiidae de la faune indo- 
malaise. Annales de la Société Entomologique de Belgique 
60: 123-136. 

Horn W., Kahlen I., Friese G., Gaedike R. 1990. Collectiones 
entomologicae. Ein Kompedium tiber den Verbleib entomo- 
logischer Sammlungen der Welt bis 1960 Teil II: L bis Z. 
Berlin: Akademie der Landwirschaften der DDR, pp. [2] + 
223-573. 

Leschen R.A.B., Löbl I. 2005. Phylogeny and classification of 
Scaphisomatini (Staphylinidae: Scaphidiinae) with notes on 
mycophagy, termitophily and functional morphology. 
Coleopterists Society Monographs, Patricia Vaurie series 
no. 3: 63 pp. 


I. Lobl 


Löbl I. 1971. Scaphidiidae von Ceylon (Coleoptera). Revue 
suisse de Zoologie 78: 937-1006. 

Löbl I. 1972. Beitrag zur Kenntnis der Scaphidiidae (Coleop- 
tera) von den Philippinen. Mitteilungen der Schweizeri- 
schen entomologischen Gesellschaft 45: 79-109. 

Löbl I. 1978. Beitrag zur Kenntnis der Gattung Sapitia Achard 
(Coleoptera, Scaphidiidae). Mitteilungen der Schweizeri- 
schen entomologischen Gesellschaft 51: 53-57. 

Lôbl I. 1981a. Über einige Arten der Gattung Scaphisoma 
Leach (Coleoptera, Scaphidiidae) aus Vietnam und Laos. 
Annales historico-naturales Musei nationalis Hungarici 
73: 105-112. 

Löbl I. 1981b. Uber die Arten-Gruppe rouyeri des Gattung Sca- 
phisoma Leach (Coleoptera Scaphidiidae). Archives des 
Sciences (Genève) 34: 153-168. 

Lôbl I. 1983. Sechs neue Scaphidiidae (Coleoptera) von 
Sulawesi, Indonesien. Mitteilungen der Schweizerischen 
entomologischen Gesellschaft 56: 285-293. 

Löbl I. 1984. Les Scaphidiidae (Coleoptera) du nord-est de 
l’Inde et du Bhoutan I. Revue suisse de Zoologie 91: 57-107. 

Lôbl I. 1986. Scaphidiidae (Coleoptera) nouveaux ou peu 
connus de l’ Asie du Sud-Est. Archives de Sciences (Genève) 
39: 87-102. 

Löbl I. 1997. Catalogue of the Scaphidiinae (Coleoptera: Sta- 
phylinidae). Instrumenta biodiversitatis 1: i-xii, 1-190. 
Löbl I. 2011. On the Scaphisomatini (Coleoptera, Staphylini- 
dae) of the Philippines, II. Revue suisse de Zoologie 118: 

695-721. 

Löbl I. 2014. On the Scaphidiinae (Coleoptera, Staphylinidae) 
of the Moluccas. Mitteilungen der Schweizerischen ento- 
mologischen Gesellschaft 87: 49-60. 

Pic M. 1915a. Diagnoses et nouveaux genres et nouvelles 
espèces de Scaphidiides. L’Echange, Revue linneene 31: 
30-32. 

Pic M. 1915b. Genres nouveaux, Espèces et Variétés nouvelles. 
Mélanges exotico-entomologiques 16: 2-13. 

Pic M. 1921. Scaphidiides recueillis par feu L. Fea. Annali del 
Museo civico di storia naturali di Genova (3) 9: 158-167. 

Pic M. 1933. Neu Coleopteren-Clavicornia. Entomologisches 
Nachrichtenblatt 7: 71-72. 


Revue suisse de Zoologie (March 2015) 122(1): 121-142 ISSN 0035-418 


Review of Brazilian cave psocids of the families Psyllipsocidae and Prionoglarididae 
(Psocodea: ‘Psocoptera’: Trogiomorpha) 
with a key to the South American species of these families 


Charles Lienhard' & Rodrigo L. Ferreira? 


! Museum d'histoire naturelle, c. p. 6434, CH-1211 Genève 6, Switzerland. Corresponding author. 
E-mail: charleslienhard@bluewin.ch 

? Universidade Federal de Lavras, Departamento de Biologia (Zoologia), CP. 3037, CEP. 37200-000 Lavras (MG), 
Brazil. E-mail: drops@dbi.ufla. br 


Abstract: Data on Brazilian cave psocids (Insecta) of the families Psyllipsocidae and Prionoglarididae are summarized, 
as a synthesis of the results of the most important investigation on cave psocids ever realized. Prionoglarididae are 
represented by 4 species of the endemic cavernicolous genus Neotrogla, Psyllipsocidae by 17 species of Psyllipsocus 
(15 of them endemic) and 2 widely distributed species of Psocathropos. These 19 recently described Brazilian endemic 
Neotrogla and Psyllipsocus were discovered in the course of the project, based on the examination of about 400 adult 
psyllipsocids and prionoglaridids collected in 124 caves situated in 59 municipalities and 13 Brazilian states. Some 
augmentations to the descriptions of the widely distributed Psyllipsocus ramburii, Psyllipsocus yucatan, Psocathropos 
lachlani and Psocathropos pilipennis are given and the following new synonymies are proposed: Psyllipsocus ramburii 
Selys-Longchamps (P. variabilis Badonnel n. syn., P dubius Badonnel n. syn.), Psyllipsocus yucatan Gurney (P collarti 
Badonnel n. syn., P decui Badonnel n. syn.), Psocathropos lachlani Ribaga (Vulturops termitorum Townsend n. syn., 
Dorypteryx astizi Brèthes n. syn.). The distribution of the 23 species of psyllipsocids and prionoglaridids known from 
Brazilian caves is analysed and some evolutionary aspects are discussed. A key to the 25 South American species of 
these families is given, including two other previously known species: Psyllipsocus delamarei from Argentina and the 
troglobitic prionoglaridid Speleopsocus chimanta from Venezuela. 


Keywords: Brazil - cave fauna - endemism - male genitalia - new synonymies. 


INTRODUCTION (Fig. 1BC) submitted to the senior author could be iden- 
tified down to species level. This material was collected 
in 124 Brazilian caves situated in 59 municipalities and 
13 states (Appendix 2). A total of 23 species belonging 
to three genera could be distinguished, 19 of them new 
to science (Appendix 1). These new species were de- 
scribed by Lienhard ef al. (2010b) and Lienhard & Fer- 
reira (2013a, b, 2014), whilst the four previously known 
non-endemic species are treated below. All psyllipsocids 
and prionoglaridids presently known from South America 
are listed (Appendix 1) and keyed. General observations 
on the distribution of the Brazilian cave species are also 
presented and some evolutionary aspects are discussed. 

This is the most important investigation on cave psocids 
ever realized (for a bibliography on cave psocids see 
Badonnel & Lienhard, 1994 and Lienhard, 2002, 2014). 
It revealed an unexpected diversity for two families 
which were almost unknown from South America, 
not only from caves but also from non-subterranean 
habitats. Some particularly interesting results of this 


All over the world, representatives of the psocid families 
Psyllipsocidae and Prionoglarididae, belonging to the 
suborder Trogiomorpha (Yoshizawa ef al., 2006), are 
regularly found in caves (Badonnel & Lienhard, 1994; 
Lienhard, 2000, 2002, 2004a, 2007; Lienhard er al. 
2010a). But both families are poorly known from South 
America (Lienhard & Smithers, 2002; Lienhard, 2004b, 
2011). We present here a synthesis of the results of a study 
of these families as they are represented in Brazilian 
caves. This study is a part of a long-term research project 
of the junior author on Brazilian cave invertebrates. 

Since 1996 some 800 caves have been visited by the ju- 
nior author and collaborating biospeleologists. Psocids of 
the above mentioned families (especially Psyllipsocidae) 
were observed in about 300 caves, mostly nymphs, but 
sometimes also adults. While the identification of the 
nymphs was not possible, the roughly 400 adult indivi- 
duals of Psyllipsocidae (Fig. 1A) and Prionoglarididae 


Manuscript accepted 26.08.2014 
DOI: 10.5281/zenodo.14579 


122 C. Lienhard & R. L. Ferreira 


Fig. 1. (A) Psyllipsocus yucatan, habitus in dorsal view (in alcohol), body length 1.5 mm (from head to abdominal tip) (after Lienhard 
et al., 2012). (B) Neotrogla curvata, habitus in dorso-lateral view (alive, in cave), body length 3.5 mm. (C) Neotrogla aurora, 
habitus in dorsal view (alive, in cave), body length 2.8 mm. 


Brazilian cave psocids 123 


study have already been presented in two special papers: 
the enigmatic presence of microcrystals on the wings of 
some specimens of Psyllipsocus yucatan by Lienhard 
et al. (2012) and the spectacular coupling role reversal 
(reversed intromittent organs) in the genus Neotrogla by 
Yoshizawa et al. (2014). 


MATERIAL AND METHODS 


The material examined was collected by RLF (unless other 
collector mentioned) by hand-collecting (for collecting 
data, see Appendix 2). Dissection and slide-mounting 
followed the methods described by Lienhard (1998). For 
general remarks and morphological abbreviations used in 
the descriptions, see Lienhard & Ferreira (2014). 

The material examined is deposited in the following 
institutions: Universidade Federal de Lavras, Depar- 
tamento de Biologia (Colecào de Invertebrados 
Subterrâneos), Lavras, Brazil (ISLA); Muséum d’ histoire 
naturelle, Geneva, Switzerland (MHNG); Systematic 
Entomology, Hokkaido University, Sapporo, Japan 
(SEHU) (some specimens of Neotrogla spp.). 
Abbreviations used in the key (nomenclature of wing 
veins according to Yoshizawa, 2005): Al = first anal 
vein; AP = areola postica (a marginal cell in forewing 
formed by veins CuAl and CuA2); CuAl = first branch 
of cubitus anterior; CuA2 = second branch of cubitus 
anterior; CuP = cubitus posterior; FW = forewing 
(length); FWw = forewing (greatest width); M = media; 
MI-M3 = branches of media; ml = medial cell anteriorly 
delimited by M1; m2 = medial cell anteriorly delimited 
by M2; P2 = second article of maxillary palp; R = radius; 
Rs = radial sector (undivided basal part of R2-R5); R1 = 
first branch of radius; rl = radial cell anteriorly delimited 
by R1; R2+3 = first branch of radial sector; r3 = radial 
cell anteriorly delimited by R2+3; R4+5 = second branch 
of radial sector; r5 = radial cell anteriorly delimited by 
R4+5; Sc = subcosta; v3 = third (external) valvula of the 
ovipositor. 

Abbreviations for Brazilian states: AL = Alagoas, AM 
= Amazonas, BA = Bahia, CE = Ceara, ES = Espirito 
Santo, GO = Goias, MG = Minas Gerais, MT = Mato 
Grosso, PI = Piaui, RN = Rio Grande do Norte, RS = Rio 
Grande do Sul, SP = Sao Paulo, TO = Tocantins. 


TAXONOMIC PART 
Non-endemic Psyllipsocidae from Brazil 


All South American species of the families Psyllipsocidae 
and Prionoglarididae are listed in Appendix 1, based on 
previously published data (Badonnel, 1962; Lienhard er 
al., 2010a, b; Lienhard & Ferreira, 2013a, b, 2014) and 
on the following data concerning four widely distributed 
Brazilian species, not treated in detail before. At present, 
these four species are the only non-endemics known 


from Brazil. The 15 endemic species of Psyllipsocus 
were treated by Lienhard & Ferreira (2013b, 2014) and 
the 4 endemic species of Neotrogla by Lienhard et al. 
(2010b), Lienhard & Ferreira (2013a) and Yoshizawa er 
al. (2014). The South American fauna of these families 
comprises 25 species, 23 of which are known from Brazil 
(Psyllipsocus delamarei only known from Argentina 
and Speleopsocus chimanta from Venezuela). See also 
the identification key for the South American species, 
presented below. 


Psyllipsocus ramburii Selys-Longchamps, 1872 
Figs 2-3 
Psyllipsocus ramburii Selys-Longchamps, 1872: 146. 
Psyllipsocus variabilis Badonnel, 1986: 182. Syn. nov. 


Psyllipsocus dubius Badonnel, 1987: 173. Syn. nov. 
For further synonymy see Lienhard & Smithers (2002). 


Material examined: ISLA and MHNG; numerous 
females, most of them micropterous (slide-mounted 
and in alcohol), collected by R. L. Ferreira in caves 
situated in the following Brazilian municipalities. — 
Castelo (ES), Gruta do Limoeiro cave, 7.1.2005. — Arcos 
(MG), Caverna do Alinhamento cave, 1.vi.2002. — 
Cordisburgo (MG), Gruta de Maquiné cave, 10.v11.2000. 
— Cordisburgo (MG), Gruta Santo Amaro 1 cave, 
28.1x.2010. — Coromandel (MG), Gruta Joao do Po 
cave, 5.x.2000. — Coromandel (MG), Gruta Ronan 
cave, 3.x.2000. — Coromandel (MG), Gruta Ronan II 
cave, 4.x.2000. — Diamantina (MG), Gruta do Salitre 
cave, 13.x11.2007. — Itumirim (MG), Gruta Santo 
Antonio cave, 25.x.2002. — Joao Pinheiro (MG), Gruta 
do Sapecado cave, 15.x.2010. — Lagoa Santa (MG), 
Gruta da Lapinha cave, 12.v1.2002. — Matozinhos (MG), 
Gruta dos Irmäos Pirias cave, 28.vii.2000. — Matozinhos 
(MG), Gruta Lavoura cave, 28.+29.v.1997, 26.v1.1997, 
19.11.2000. — Matozinhos (MG), Meandro Abismante 
cave, 6.vii.2002. — Matutina (MG), Gruta 9 cave, 
10.x.2010. — Moeda (MG), cave SMS 29, 11.x11.2005. 
— Pains (MG), Gruta do Capäo cave, 5.v.2001. — Pains 
(MG), Gruta Paranoa cave, 15.1.2008. — Pains (MG), 
Gruta Retiro cave, 27.xi.1999. — Paracatu (MG), Gruta 
da Fazenda Tamandua II cave, 14.x.2010. — Paracatu 
(MG), Lapa do Brocoté cave, 16.1x.2010. — Paracatu 
(MG), Lapa de Santo Antonio cave, 13.vi1.2010. — Santa 
Maria do Suaçui (MG), Gruta do Rio Suaçui cave, 
19.vii.2002. — Sete Lagoas (MG), Gruta Rei do Mato 
cave, 3.+4.xi.2011. — Vazante (MG), Lapa da Delza 
cave, 12.vii.2010. — Torres (RS), Furna da Lagoa de 
Itapeva cave, 20.v.2008. — Altinöpolis (SP), Gruta do 
Itambé cave, 27.11.2006. — Altinöpolis (SP), Gruta Olho 
de Cabra cave, 2.111.2006. — Altinöpolis (SP), Gruta do 
Parana cave, 1.i11.2006. — Itirapina (SP), Gruta da Toca 
cave, 22.x.2004. 


Further description: P. ramburii is the type species 
of the genus Psyllipsocus; it is polymorphic and 
occurs in three forms: macropterous, brachypterous 


124 C. Lienhard & R. L. Ferreira 


ES N 
| 


Fig. 2. Psyllipsocus ramburii, from Brazilian caves, macropterous female MHNG 7929 from Gruta Santo Antônio (A-D, H), 
micropterous female MHNG 7954 from Gruta do Rio Suaçui (E-G). (A) Forewing. (B) Hindwing. (C) P2-P4 of maxillary 
palp. (D) P2-chaetotaxy. (E) Habitus (body length 1.7 mm). (F) Pedicel, showing absence of microspades organ (compare with 
Fig. 7D). (G) Spermapore plate and spermatheca. (H) Spermatheca. 


Brazilian cave psocids 125 


> DS 


TIP 


SK 


Fig. 3. Psyllipsocus ramburii, macropterous female (MHNG 8125) with slightly shortened wings, from Gruta Joao do P6 cave (Brazil, 
MG, Coromandel). Forewings and hindwings (pilosity not shown). 


and micropterous; the existence of an apterous form is 
somewhat doubtful and has yet to be confirmed (see 
Key and Discussion, below). For detailed descriptions 
see Mockford (1993) and Lienhard (1998). Two 
characters are also especially noted by Mockford 
(2011): “macropterous form lacking distal closed cell 
in forewing; setal organ at base of paraproctal spine 
consisting of one very long and one short seta.” Here 
we note the following additional characters (Figs 2-3, 
see also Key): pedicel lacking microspades organ (Fig. 
2F); P2 lacking stout sensillum (Fig. 2D); spermatheca 
small and crumpled (due to parthenogenesis), with 3-4 
small and irregularly shaped sclerites near duct (Fig. 
2GH; see also Lienhard, 1998: fig. 37e and Badonnel, 
1986: fig. 22); spermapore region with a characteristic 
pair of more or less sclerotized longitudinal straps (Fig. 
2G; see also Mockford, 1993: fig. 62 and Badonnel, 
1986: fig. 22); macropterous form occasionally with 
slightly shortened wings (not as short as in the typical 
brachypterous form, see Lienhard, 1998: figs 37c, 38c) 
and rather variable venation (Fig. 3, see also Badonnel, 
1986: figs 14-18); in micropterous form length of 
minute wing rudiments variable, relatively long and 
bearing some vein rudiments (Lienhard, 1998: fig. 37a, 
g) or very short and veinless (Fig. 2E). 


Biology: P ramburii is a parthenogenetic species 
(thelytoky) and only one single (accidental?) male is 
known (Lienhard, 1977, 1998). The phallosome of the 
male is equipped with a pair of well-developed basal 
struts, as they are present in all psyllipsocids (Mockford, 
2011) except for most of the Brazilian cave endemics 
(Lienhard & Ferreira, 2014; see also Evolutionary 
Considerations, below). 


Distribution: P ramburii is a widespread species 
(Lienhard & Smithers, 2002), often domicole but also 
regularly found in caves all over the world (Badonnel & 


Lienhard, 1994). Previously published South American 
records can be found in the following references: 
Strinati, 1971 (Uruguay); Badonnel, 1972 (Chile); 
Badonnel, 1986 (Colombia, as P variabilis); Badonnel, 
1987 (Venezuela, as P dubius); Cammousseight & New, 
1994 (Chile). 


Discussion: The presence, in the macropterous 
specimens from Brazil, of a variability of wing venation 
similar to that described by Badonnel (1986) for 
P. variabilis from Colombia, and the absence of other 
distinctive characteristics, support synonymizing the 
latter with P ramburii. 

The case of P dubius is more complex. This apterous 
species was described from Venezuela by Badonnel 
(1987), based on the holotype female from cacao ground 
litter and a second (non-paratype) female from a cave. 
The microscopical slides of both specimens, mounted by 
André Badonnel himself, are deposited in the MHNG and 
were examined. Unfortunately the thorax of the holotype 
had been damaged and only the prothorax is present on 
the slide. Thus, the absence of wing rudiments could not 
be confirmed for this specimen. The second specimen 
is better preserved; it is complete, though its thorax 
was deformed by slide-mounting. Surprisingly it bears 
wing rudiments which are very similar to those of the 
micropterous form of P ramburii; in this female, they are 
particularly small and completely veinless (asin the female 
from a Brazilian cave figured in Fig. 2E). This specimen 
of P. dubius has clearly to be assigned to P ramburii. In 
the original description of P dubius, Badonnel insists 
on the complete apterism of this species and does not 
mention any variability of wing development. He also 
mentions that the only morphological difference between 
P ramburii and P. dubius is the complete absence of wing 
rudiments in the latter. Even though the holotype might 
really be apterous, we are convinced that synonymy of 
P. dubius With P ramburii is justified. The terminalia 


126 C. Lienhard & R. L. Ferreira 


of the holotype correspond exactly to P ramburii; in 
particular the spermapore region with the typical pair of 
sclerotized longitudinal straps clearly visible. However, 
the quite plausible existence of an apterous form of this 
polymorphic species is not yet definitively confirmed 
with this synonymization. It is possible that Badonnel 
did not realize that the part of the broken thorax of the 
holotype present on the slide was only the prothorax and 
misinterpreted the absence of wing rudiments on this part 
as apterism, while he probably overlooked the small wing 
rudiments present on the deformed slide-mounted thorax 
of the second female. Alternatively, it is also possible that 
he had already observed the absence of wing rudiments in 
the holotype before dissecting the specimen; impressed 
by this unusual character he might have overlooked their 
presence in the second female. 


Psyllipsocus yucatan Gurney, 1943 
Figs 1 A and 4-6 


Psyllipsocus yucatan Gurney, 1943: 212. 
Psyllipsocus collarti Badonnel, 1946: 140. Syn. nov. 
Psyllipsocus decui Badonnel, 1977: 340. Syn. nov. 


Material examined: ISLA and MHNG; numerous 
males and females (slide-mounted and in alcohol), 
collected by R. L. Ferreira (unless other collector 
mentioned) in caves situated in the following Brazilian 
municipalities. — Murici (AL), Toca da Raposa | cave 
(granite), 13.1.2007. — Murici (AL), Toca da Raposa 2 
cave (granite), 13.1.2007. — Campo Formoso (BA), Toca 
da Tiquara cave, 8.1.2008, 1.2009, x11.2010. — Santa 
Maria da Vitoria (BA), cave PEA 343, 15.v.2011, leg. 
S. S. Salgado. — Ubajara (CE), Gruta dos Mocös cave, 
3.1.2007. — Ubajara (CE), Gruta do Morcego Branco 
cave, 3.1.2007. — Ubajara (CE), Gruta de Ubajara 
cave, 30.x11.2006. — Castelo (ES), Gruta do Limoeiro 
cave, 7.1.2005. — Arinos (MG), Lapa do Salobo cave, 
18.v11.2010. — Santa Maria do Suaçui (MG), Gruta do 
Rio Suaçui cave, 19.v11.2002. — Teofilo Otoni (MG), 
Lapa da Vaca Parida cave, 26.1.2005. — Felipe Guerra 
(RN), Caverna Trapia cave, 6.1.2010, 4.viii.2010, leg. 
D. M. Bento. — Ilha Bela (SP), Gruta da Serraria cave, 
22.1v.2006. 


Further description: This species is always macro- 
pterous. For detailed descriptions see Gurney (1943) 
and the augmented description given by Mockford 
(2011). Here we note the following characters (Figs 4-6, 
see also Key). Forewing usually with a subtle but typ- 
ical colouration (Fig. 4G), as described by Gurney 
(1943: p. 212): “membrane faintly tinged with fuscous, 
a clear spot at posterior margin of wing between apex 
of Cu2 [= CuA2] and apices of anal veins [= CuP and 
Al].” P2 lacking clearly differentiated stout sensil- 
lum (Fig. 4D). Epiproct characteristically pigment- 
ed (Fig. 4BF). Paraproctal setal organ consisting of 
a minute hair and a longer and thicker seta; anal spine 


replaced by a stout, relatively short and basally arti- 
culated seta (not much longer than long seta of setal 
organ) (Fig. 4BF); female paraproct simple, male para- 
proct ventrally with a small concave protuberance 
(“fossette inférieure” of Badonnel, 1946) (Fig. 4F). 
Pigmentation of hypandrium interrupted in middle of 
anterior margin (Fig. 6AB); aspect of distal phallosome 
sclerites somewhat variable, depending on view (dorsal/ 
ventral) and on position after slide-mounting (Fig. 6A 
similar to fig. 5 in Badonnel, 1946; Fig. 6B similar to 
fig. 107 in Mockford, 2011). Sclerotization of oviposi- 
tor vl and v2 variable; sclerotized median axis of vl 
weakly developed or absent; median axis of v2 usual- 
ly well-sclerotized (Fig. 5A), but weakly developed 
or completely absent in pale individuals (so observed 
in several pale specimens from Brazil, Antigua and 
Thailand). Spermapore plate weakly sclerotized but 
characteristic (Fig. 5B); spermathecal sac with complex 
sclerotizations near origin of duct (Fig. SDE); spermato- 
phores not sclerotized, somewhat variable in shape, but 
always similar to that figured in Fig. 5C; the spermathe- 
ca of a female may contain up to about a dozen sper- 
matophores (see legend to Fig. 5B). 


Biology: In general, P yucatan seems to be a cave 
species and it was explicitly indicated as living on 
vegetation only once (Badonnel, 1948: on leaves of an 
orange tree), with no domicole population known. The 
presence of up to about a dozen spermatophores in the 
spermatheca of one single female indicates that the 
species is polyandrous. 


Distribution: P yucatan was originally described from 
a Mexican cave by Gurney (1943). Previously published 
South American records can be found in the following 
references: Garcia Aldrete & Mockford, 2009 (Brazil, 
probably same record as that mentioned by Mockford, 
2011); Mockford, 2011 (Brazil, in an artificial cave); 
Lienhard ef al., 2012 (Brazil, in caves). The species is 
also known from a cave in Cuba (Badonnel, 1977, as 
P. decui) and from caves in Jamaica, Aruba and Antigua 
(MHNG, unpublished). African specimens are known 
from Congo (Badonnel, 1946, 1948, as P collarti, on 
vegetation), Kenya and Senegal (MHNG, unpublished). 
Some specimens of P yucatan have also been collected 
in a cave in Thailand (MHNG, unpublished). The 
species seems to be widely distributed in the tropics. 


Discussion: The only difference between P. yucatan and 
P. decui indicated by Mockford (2011) is the absence of 
the sclerotized median axis of v2. The above mentioned 
variability shows that this character cannot be used 
to separate these species. The absence of a sclerotized 
v2-axis in the palest specimens examined corresponds 
to the situation described by Badonnel (1977) for the 
very pale holotype of P decui from Cuba, where vl and 
v2 are completely hyaline. Therefore P. decui is here 
considered as a junior synonym of P yucatan. 


Brazilian cave psocids 127 


Fig. 4. Psyllipsocus yucatan, from Brazilian caves, general morphology. (A) Habitus, male. (B) Epiproct and right paraproct, female. 
(C) Lacinial tip, female. (D) P2-chaetotaxy, female. (E) P2-P4 of maxillary palp, female. (F) Epiproct and left paraproct, male. 
(G) Forewing, male. (H) Hindwing, male. 


128 C. Lienhard & R. L. Ferreira 


aN 


ES 


N 


Fig. 5. Psyllipsocus yucatan, from Brazilian caves, female genitalia. (A) Subgenital plate, right ovipositor valvulae and right hind 
corner of clunium. (B) Spermapore plate and spermatheca (the latter contains about a dozen spermatophores, only one of them 
shown). (C) Spermatophore (not same female as B). (D) Spermathecal sclerotizations and proximal part of spermathecal duct, 
specimen MHNG 7990 from Toca da Raposa 2, AL. (E) Ditto, specimen MHNG 8000 from Gruta de Ubajara, CE. 


Brazilian cave psocids 129 


Fig. 6. Psyllipsocus yucatan, from Brazilian caves, male genitalia. (A) Hypandrium and phallosome, ventral view. (B) Ditto, dorsal 
view, other specimen than in A (hypandrium schematically shown by interrupted lines). 


Badonnel (1948, 1977) and Mockford (2011) noted the 
close proximity of P yucatan and the African P. collarti. 
CL has examined the series of syntypes of P collarti 
(1 female and 1 male slide-mounted by Badonnel, the 
latter on two separate slides, and several specimens in 
alcohol, some of them badly damaged, all deposited 
in the Musée Royal de l’Afrique Centrale, Tervuren, 
Belgium). No significant difference between the African 


and the Brazilian specimens could be observed. The 
structure figured by Badonnel (1946: fig. 10, labeled vv) 
as a sclerotized distal appendage of vl is only visible on 
one side of the slide-mounted terminalia; it is evident that 
this is an artefact (probably a particle of detritus adhering 
to the membranous v1). P. collarti is here considered as a 
junior synonym of P yucatan. 


130 C. Lienhard & R. L. Ferreira 


Psocathropos lachlani Ribaga, 1899 
Fig. 7 


Psocathropos lachlani Ribaga, 1899: 157. 

Vulturops termitorum Townsend, 1912: 269 (type species of 
Vulturops). Syn. nov. 

Psocathropos termitorum (Townsend); Roesler, 1944: 
134 (Vulturops considered as a junior synonym of 
Psocathropos). 

Dorypteryx astizi Brèthes, 1923: 117. Syn. nov. 

Psocathropos astizi (Bréthes); Lienhard, 1977: 435 (new 
combination suggested). 

Psocathropos astizi (Brèthes); Lienhard & Smithers, 2002: 43. 

For further synonymy see Lienhard & Smithers (2002). 


Material examined: ISLA and MHNG; several males 
and females (slide-mounted and in alcohol), collected by 
R. L. Ferreira in caves situated in the following Brazilian 
municipalities. — Delmiro Gouveia (AL), Gruta Morcego 
cave, 23.iv.2006. — Curaçä (Patamuté) (BA), Toca d’agua 
de Patamuté cave, 6.1.2008. — Ubajara (CE), Gruta de 
Ubajara cave, 30.x11.2006. — Santa Teresa (ES), Gruta 
do André Huscki cave, 4.1.2005. — Cordisburgo (MG), 
Gruta de Maquiné cave, 10.vi1.2000 and 18.v.2010. — 
Montalvania (MG), Gruta Nossa Senhora do Perpétuo 
Socorro cave, 14.v11.2007. — Nacip Raydan (MG), Gruta 
Manga de Pedra cave (granito), 21.v11.2002. — Teofilo 
Otoni (MG), Lapa da Vaca Parida cave, 26.1.2005. — 
Altinépolis (SP), Gruta do Itambé cave, 27.11.2006. 


Further description: P /achlani is the type species of 
the genus Psocathropos; it is usually brachypterous. The 
wings of the rare macropterous form (see Key) were 
first described by Mockford (1993); two macropterous 
females from a laboratory culture (Jamaica: Kingston, 
leg. B. D. Turner, 1971) are deposited in the MHNG. 
For detailed descriptions see Mockford (1993) and 
Lienhard (1998); some important characters are also 
mentioned by Badonnel (1967, see Discussion below) 
and in the key given by Mockford (1991). Here we note 
the following characters (Fig. 7, see also Key). Length 
of forewing of brachypterous form variable, usually 
not reaching abdominal tip; venation of forewing also 
somewhat variable, but Rs always simple (Fig. 7A). 
Hindwing of brachypterous form always reduced to a 
minute bare and veinless flap (Fig. 7B). P2 lacking stout 
sensillum. Pedicel with well-developed microspades 
organ (Fig. 7E). Paraproctal trichobothria with weakly 
differentiated basal florets; setal organ consisting of one 
minute hair and one somewhat thicker and longer seta; 
anal spine not differentiated, replaced by a normal seta 
which is not much longer than the ventral seta of the 
setal organ (Fig. 7FG). Epiproct and paraproct of female 
with some conspicuous cylinder setae (Fig. 7F). Setae 
of male epiproct and paraproct normal, but paraproct 
ventrally pointed due to the presence of a conspicuous 
posteroventral sclerotized ridge which is ventrally 
linked to fine membranes (Fig. 7G); depending on slide- 
mounting this structure may appear like a free-ending 


appendix (see Badonnel, 1967: fig. 58; Mockford, 
1993: fig. 467; Lienhard, 1998: fig. 36j). Hypandrium 
and phallosome as in Fig. 7H, hypandrium shorter 
than its basal width, phallic cradle well differentiated; 
longitudinal struts in middle connected to each other by 
a V-shaped sclerite, pores on apical part of longitudinal 
struts present but not visible in Fig. 7H (only visible 
in strongly squashed genitalia; see Badonnel, 1967: 
fig. 57). Spermatheca as in Fig. 7D, with a sclerotized 
globular vesicle near duct, spermapore lacking 
sclerotization, spermathecal sac with a narrowed zone 
towards opening of duct, spermatophore about pear- 
shaped. 


Biology: In relatively warm regions, P. lachlani is 
often domicole, as was the type population from 
southern Italy (Ribaga, 1899); in the tropics it is also 
regularly found in ground litter and sometimes in 
caves (Lienhard, 2002). Bréthes (1923) and Townsend 
(1912) recorded the species indoors (as P. astizi and 
P. termitorum respectively; see Discussion, below), the 
latter especially in houses infested with termites. The 
presence of one “fresh” spermatophore and debris of 
about three “old” spermatophores in the female figured 
(Fig. 7D) indicates that the species is polyandrous. 


Distribution: P. lachlani is widespread (Lienhard & 
Smithers, 2002). Previously published South American 
records can be found in the following references: 
Townsend, 1912 (Peru, as P termitorum); Brèthes, 
1923 (Argentina, as P astizi); New, 1984 (Brazil); 
Mockford, 1993 (Colombia); Garcia Aldrete & 
Mockford, 2009 (Brazil). In adddition to the Brazilian 
specimens mentioned above, some unpublished 
material from Paraguay (MHNG) was also examined. 
Other specimens examined (MHNG) are from the 
following countries: Guatemala, Guadeloupe, Jamaica, 
Ascension Island, Cabo Verde, Madeira, Liberia, Togo, 
Rwanda, Mozambique, Morocco, Israel, Reunion, India, 
Malaysia, Singapore, Indonesia, Thailand, Fiji. 


Discussion: The detailed descriptions and the forewing 
figures given by Townsend (1912) and Brèthes (1923) 
for P termitorum and P. astizi respectively fit very well 
the widely distributed P /achlani; therefore we propose 
here to consider these two names as junior synonyms of 
the latter. 

Under the synonym P microps (Enderlein), Badonnel 
(1967: p. 27) gives excellent figures of the forewing 
and the male terminalia of P lachlani, but it has to be 
noted that in the legend (not in the text!) his figure 58 is 
erroneously assigned to P. pilipennis. 


Psocathropos pilipennis (Enderlein, 1931) 
Fig. 8 


Gambrella pilipennis Enderlein, 1931: 221. 
Psocathropos pilipennis (Enderlein); Menon, 1942: 35. 


Brazilian cave psocids 131 


Fig. 7. Psocathropos lachlani, from Brazilian caves. (A) Forewing, female. (B) Hindwing, female. (C) Left ovipositor valvulae and 
left hind corner of clunium. (D) Spermatheca containing one “fresh” spermatophore and debris of three “old” spermatophores. 
(E) Pedicel with microspades organ, female. (F) Right paraproct, female. (G) Epiproct and paraprocts, male. (H) Hypandrium 
and phallosome, ventral view (pilosity not shown). 


132 C. Lienhard & R. L. Ferreira 


Fig. 8. Psocathropos pilipennis, from Brazilian caves. (A) Forewing, male (pilosity not shown). (B) Hindwing, male, showing venation 
and insertion points of setae. (C) Right paraproct, female. (D) Spermathecal sclerotizations and proximal part of spermathecal 
duct, specimen MHNG 8148 from Toca do Garrincho, PI. (E) Spermatheca (duct incomplete) containing one spermatophore, 
specimen MNHG 8150 from Gruta do Araticum, CE (sclerotizations less squashed than in D). (F) Hypandrium and phallosome, 


ventral view (pilosity not shown). 


Material examined: ISLA and MHNG; 4 males and 
4 females (slide-mounted and in alcohol), collected 
by R. L. Ferreira in caves situated in the following 
Brazilian municipalities. — Ubajara (CE), Gruta do 
Araticum cave, 1.1.2007. — Coronel José Dias (PI), Toca 
do Garrincho cave, 11.ix.2008. — Felipe Guerra (RN), 
Gruta Roncador cave, 1.vi.2006. — Jandaira (RN), Gruta 
Aroeira cave, 11.1.2006. 


Further description: This species is always 
brachypterous. For general descriptions see Enderlein 


(1931: female) and Badonnel (1967: male, see also 
Discussion below); some important characters are also 
mentioned in the key given by Mockford (1991) and 
in Mockford (1993: p. 51). Here we note the following 
characters (Fig. 8, see also Key). Body length 1.4- 
1.6 mm. Colouration similar to P lachlani, but usually 
less pigmented, sometimes hypodermal pigment almost 
absent. Forewing slightly tapering towards apex, usually 
almost reaching tip of abdomen or slightly surpassing it 
(especially in males); venation of forewing somewhat 


Brazilian cave psocids 13 


variable, but Rs always bifurcate (Fig. 8A). Hindwing 
strongly shortened but always with some clearly 
differentiated veins and several long marginal setae 
(Fig. 8B). P2 and pedicel as in P lachlani. Epiproct and 
paraproct simple in both sexes (i.e. no cylinder setae in 
female and no paraproctal posteroventral ridge in male), 
anal spine replaced by a normal seta, setal organ as in 
P. lachlani (Fig. 8C). Hypandrium and phallosome as in 
Fig. 8F, hypandrium longer than its basal width, apically 
truncate, phallic cradle not clearly differentiated; 
longitudinal struts lacking V-shaped connecting sclerite, 
apically with clearly visible pores. Spermatheca as in 
Fig. 8E, sclerotizations near duct more complicated than 
in P. lachlani and vesicle slightly elongate (Fig. SDE). 
Spermapore lacking sclerotization, spermathecal sac 
oval, lacking narrowed zone towards opening of duct, 
spermatophore elongately pear-shaped (Fig. 8E). 


Biology: In houses (Menon, 1942; Badonnel, 1967; 
Mockford, 1991), in ground litter and on low vegetation 
(New, 1977), in caves. 


Distribution: Published records are from the Seychelles 
(Enderlein, 1931), India (Menon, 1942), Madagascar 
(Badonnel, 1967) and the Aldabra Islands (New, 1977). 
In addition to the Brazilian material, some specimens 
from the following countries have also been examined 
(MHNG): Honduras (in cave), Curacao (in cave), 
Singapore (ground litter and/or in houses), Malaysia 


(99) 


(on fungi in forest), Vietnam (ground litter), Thailand 
(domicole?). The species seems to be widespread 
in the tropics, though less common than P lachlani. 
However, it is not excluded that some published records 
of the latter comprise also undiscovered material of 
P. pilipennis (e.g. Menon, 1942; see Discussion below). 
Badonnel (1967) also indicates that both species were 
taken together in a house in Madagascar. 


Discussion: Menon (1942) synonymizes the genus 
Gambrella Enderlein with Psocathropos Ribaga and 
gives a description of its type species, P. pilipennis. 
However, his material apparently consisted of a 
mixture of both species, P pilipennis and P lachlani. 
Therefore he mentions a variability of wing morphology 
(hindwing sometimes “totally wanting” and forewing 
sometimes relatively short with simple Rs, see figures 
25-26) which we have never observed in P pilipennis. 
In our opinion, his figures 25 and 26 illustrate forewings 
of P lachlani, while his figures 23, 24 and 27 illustrate 
wings of P. pilipennis, and his figure 28 the male 
genitalia of the latter. Badonnel (1967) gives excellent 
figures of the wings and the male genitalia, but it has to 
be noted that his figure 58 does not illustrate epiproct 
and paraproct of P pilipennis, as erroneously indicated 
in the legend, but of P /achlani (= P. microps), as 
correctly indicated in the text on the latter species 
(pe): 


Key to the South American species of Psyllipsocidae and Prionoglarididae 


This key allows species identification of adult South American psocids belonging to the families Psyllipsocidae and 
Prionoglarididae. In addition to the characters mentioned in the key, these species are characterized among Psocoptera 
by their 3-segmented tarsi, the absence of scales on body and wings, the absence of a thickened and distinctly opaque 
pterostigma (sometimes slightly opaque in Prionoglarididae) and the presence, in the forewing, of a simple anal vein and 
a nodulus (i.e. veins CuP and Al joining at wing margin). Their body is never dorsoventrally flattened and, if present, their 
forewings are membranous, never coriaceous, though sometimes slightly vaulted (elytriform). 

For abbreviations see Material and Methods, above. 


Forewing normally developed (macropterous) or slightly shortened, usually extending well beyond tip of abdomen, 
not tapering towards apex but broadly rounded, with normal venation …............................................................ 2 
Apterous, micropterous or clearly brachypterous (forewing usually not reaching tip of abdomen or tapering towards 
APEX)AVENATIONANIEAS SIC NTIVÉTEQUC el RS Nn re 3 
Forewing: basal portion of Sc developed as a slightly curved vein joining R1 just basally of pterostigma (Lienhard 
Sakerreiraw OS asi casita) SS odyzlensth eon OPS AMIE seer eee eee Prionoglarididae: Neotrogla 11 
Forewing: basal portion of Sc rudimentary, ending in membrane, not joining R and usually not extending beyond 
base of basal closed cell (Fig. 2A). Body length: 1.0 mm (Psyllipsocus didymus) to 2.7 mm (P. serrifer), mostly 
ADOULEES S00 IMPERO RR e e TE AT ta el La Psyllipsocidae 14 
APLELOUS MWithOUtMWin LIU diMeNTSALOCENTTEOT PIELE VARE (LG E de 4 
NTICTO PLIETO USTOTIDEACHYPLETO USAI E Re en A TR 5 
Pretarsal claw without preapical tooth. Eye minute, consisting of a single ommatidium. Paraproct without anal 
STINE PR cee svar cee eee IT se PEUR CE M RUE IA tei Prionoglarididae: Speleopsocus chimanta 
Note: This species is a troglobite known from a cave in south eastern Venezuela (see Lienhard er al., 2010a). 

Pretarsal claw with a small preapical tooth. Eye differentiated as a small compound eye, with several ommatidia. 
Raraproctawithtaslonoramalis DIN En sacusea ccosusesseesesectesss en Psyllipsocus ramburii 


C. Lienhard & R. L. Ferreira 


Note: This is the rare (or potential?) apterous form of P ramburii, corresponding to P dubius Badonnel which 
is here considered as a junior synonym of the latter (see Discussion of P ramburii, above). For micropterous, 
brachypterous and macropterous forms, see couplets 5, 10 and 15. 

Forewing flat or slightly vaulted (elytriform), reaching posterior half of abdomen and bearing some clearly 


Gifferentiated: VEINS 2287 huh ae ee Se RE ae RE RE 6 
Forewing strongly reduced, just reaching basal abdominal segments and lacking clearly differentiated veins 
(Bisa B) SRaraproctawithyaslongganiallls pin CE Psyllipsocus ramburii 


Note: This is the micropterous form of P ramburii; for apterous, brachypterous and macropterous forms, see 
couplets 4, 10 and 15. 
Forewing margin with long setae (their length several times width of marginal vein) (Fig. 7A). Paraproct lacking 


a cOnspicuousranalespine (HiSSi 7K GRO) cere REA eee Psocathropos 7 
Forewing margin bare (except for some microscopic hairs which are much shorter than width of marginal vein). 
Paraproct with a long anal spine (Lienhard & Ferreira, 2014: fig. 1F)........... Psyllipsocus 8 


Forewing Rs simple (Fig. 7A); hindwing strongly reduced, bare and veinless (Fig. 7B). Hypandrium shorter than 
its basal width (Fig. 7H). Male paraproct ventrally pointed, with a posteroventral sclerotized ridge (Fig. 7G). 
Female paraproct and epiproct with some conspicuous cylinder setae (Fig. 7F) .................. Psocathropos lachlani 
Note: This is the brachypterous form of P lachlani; for the rare macropterous form, see couplet 14. 

Forewing Rs bifurcate (Fig. 8A); hindwing much shortened but with some clearly differentiated veins and some 
long marginal setae (Fig. 8B). Hypandrium longer than its basal width (Fig. 8F). Male paraproct as in female 
(Fig. 8C), ventrally rounded, lacking sclerotized ridge. Female paraproct and epiproct lacking cylinder setae ....... 
TEE Er“ Psocathropos pilipennis 
Note: Macropterous form unknown in this species. 

Forewing with a brown transversal band about in middle (Lienhard & Ferreira, 2014: fig. 1C)........... n 


EEE TR IO A RE ET AR Psyllipsocus spinifer 
Note: For the much rarer macropterous form of this species, see couplet 22. 

Eorewinp:unmarked....-.- rr elio n 9 
P2 with a stout sensillum in basal half (Lienhard & Ferreira, 2013b: fig. 4F).................... Psyllipsocus clunjunctus 
Note: Macropterous form of this species not known. 

StouwP2=sensillummotfdifferentiated{(B192) FR OI 10 
Forewing slightly vaulted (elytriform), hindwing a tiny membranous flap only reaching hind margin of first 
abdominal tergite? =. GERMI TOA Psyllipsocus delamarei 


Note: This species is only known from one female from Argentina (see Badonnel, 1962); it seems to be very close 
to the brachypterous form of P ramburii. 

Forewing not elytriform, hindwing with some veins, not much shorter than half length of forewing ...................... 
QUORE RAA M RR, e RER NERRNSOER RSERREHEEN Psyllipsocus ramburii 
Note: This is the brachypterous form of P ramburii; for apterous, micropterous and macropterous forms, see 
couplets 4, 5 and 15. 


Female: distal part of gynosome strongly curved (Lienhard & Ferreira, 2013a: fig. 1d).............. Neotrogla curvata 
Female: distal part of gynosome straight or only slightly curved (Lienhard er al., 2010b: figs 2df, 8c) ............... 12 
Female: membranous part of gynosome not lobate (Lienhard er al., 2010b: fig. 8C).................. Neotrogla truncata 
Female: membranous part of gynosome with a spiny dorsal lobe and a pair of spiny lateral lobes (Lienhard er al., 
20106: fies DL) is. set EN LEZIC RIZZI RI RE CREO 13 
Female: distal lobe of subgenital plate (egg-guide) apically with a large bare median lobe and a pair of minute 
pilosellateralMobesi((&Tenhardieia/Z2010b4f A) Neotrogla brasiliensis 


Female: pilose lateral lobes of egg-guide not much smaller than bare median lobe (Lienhard ef al., 2010b: fig. 5) 

vale ella i ea Eee SIERRA CREA Re Neotrogla aurora 
Forewing and hindwing with numerous long marginal setae (their length several times width of marginal vein) 
(Mockford;‘1993=:f1ps:462£ 463) 2.2.2... EE TO Psocathropos lachlani 
Note: This is the rare macropterous form of P lachlani; for the brachypterous form, see couplet 7. 

Wing margins bare (except for some microscopic hairs which are shorter than width of marginal vein)............. 15 
Distal closed cell in forewing absent (i.e. no crossvein between RI and Rs) (Fig. 2A) ......... Psyllipsocus ramburii 
Note: This is the relatively rare macropterous form of P ramburii; for apterous, micropterous and brachypterous 
forms, see couplets 4, 5 and 10. 


Distal closed cell present, distally delimited by a crossvein between RI and Rs (Fig. 4G) ...... in 16 
In forewing Rs and M joined by a crossvein (Lienhard & Ferreira, 2014: fig. 3A).............................................. 17 
InkforewingiRsfandiMéfusedkforgallength{(BIZ4) rene 19 


Forewing.unmarked'..:.....inrca dB EI O Psyllipsocus falcifer 


22 


25 


24 


2S 


26 


DA] 


28 


29 


Brazilian cave psocids 135 


Korewingswithtsomespismentedfare AS nn nn ceci ine sense sie ennui e ns sans anni ee 18 
AP flat, its marginal length more than twice its height, patches of wing pattern associated to veins (Lienhard & 
Retrelra 204 OSARE RE onori coca sees soctedocs sossecsisscsdstencedssehacsatcncesocevedesrsee Psyllipsocus marconii 
AP tall, its marginal length not much exceeding its height, an isolated longitudinal patch present in middle of cells 
ell, 103} eral eS) (Lieiolnereal 6%, Merman, OCEAN Psyllipsocus thaidis 
First portion of pterostigmal R1 not longer than half length of RI-Rs crossvein (Fig. 4G) .............. n 20 
First portion of pterostigmal RI clearly longer than half length of R1-Rs crossvein, often longer than this 
ERINNERN ohne n NR nai 22) 
Distal closed cell almost twice as long as marginal length of pterostigma; paraproct with a long, basally non- 
arttculatedtanallspinef(ikienhardreKerreira2 OS ba SSH en nine enr SITO 21 
Distal closed cell only about half as long as marginal length of pterostigma (Fig. 4G); anal spine of paraproct 
replaced by a stout, relatively short and basally articulated seta (Fig. 4BF)............................ Psyllipsocus yucatan 


Female: spermatheca with a conspicuous longitudinal saw-like sclerite (Lienhard & Ferreira, 2013b: fig. 7J). Male: 
abdominal apex with a complex clunial bridge, hypandrium with four apical setae (Lienhard & Ferreira, 2013b: 
LI PRI) Re en eee ee eee eee re eee Psyllipsocus serrifer 
Female: spermatheca with a slender file-like sclerite (Lienhard & Ferreira, 2013b: fig. 8F). Male: abdominal apex 
with a simple clunial bridge, hypandrium with two apical setae (Lienhard & Ferreira, 2013b: fig. 9) ..................... 
GED GORE DROLE SONO AEE RATTI a Psyllipsocus similis 
Forewing and hindwing with a brown transversal band in basal half (Lienhard & Ferreira, 2014: fig. 1 AB) .......... 
RR CRON oat nia la iena Psyllipsocus spinifer 
Note: This is the relatively rare macropterous form of this species; for the brachypterous form, see couplet 8. 


WINE stunmarkedfor@win IP ALLIE CT EN Er 25 
Rorewinegpanticu laniygn ATOME EN ES TR n nr 24 
Hore win 210 normal ha DEMAIN ES PR tn n OOOOH re nent eee e 25 


Forewing with narrow pigment bands bordering most of veins (Lienhard & Ferreira, 2014: fig. 16A). Male: 
postero-median V-shaped incision of phallosome not deeper than length of antero-median part of phallosome 
selenite (Ligal AT 6s emarmga, IPN CAIN) ee Psyllipsocus angustipennis 
Pigment bands broader in apical half of forewing, covering more than half of the surface of cells ml and m2 
(Lienhard & Ferreira, 2014: fig. 17C). Male: postero-median V-shaped incision of phallosome deeper than length 
of antero-median part of phallosome sclerite (Lienhard & Ferreira, 2014: fig. 17F)............. Psyllipsocus proximus 
Forewing anal vein with a brown spot at the base of each hair; exceptionally forewings completely unmarked (see 
Paleispecimensio PRES TD EH IIS ACOUD ES) PR I O TRO trio 26 
Forewing anal vein lacking such spots; distal end of forewing veins bordered by brown pigment (Lienhard & 
Rérretra 04356 ES TANIA) Psyllipsocus punctulatus (and female spec. cf. punctulatus) 
Height of AP exceeding its marginal length; forewing with a conspicuous U-shaped or V-shaped brown patch 
covernpimosttoffcellin1t(Ienhardi&AFerreira 2014 figs WS A) ee Psyllipsocus radiopictus 
AEglowerthangtsimareinallength34piomenta o No NcerNditfere NE 27 
Forewing with a broad brown band bordering apical portion of R1 and RI-Rs crossvein, forming a conspicuous 
angulate patch which borders cell rl basally (Lienhard & Ferreira, 2014: fig. 12A)............................................... 
MRI IRR RIINA IIS SHORES EE SII PROT AA Psyllipsocus fuscistigma 
At most narrow pigment bands bordering apical portion of RI and RI-Rs crossvein..................... i 28 
Forewing with four narrow pigment stripes parallel to veins R2+3 and R4+5, situated in cells rl (one stripe), r3 
(two stripes) and r5 (one stripe); these stripes clearly separated from the narrow pigmented zones bordering these 
veins. In very pale specimens forewings unmarked. Male: clunium simple, lacking clunial rods. Female: v3 with 
a marginal row of thick setae, clearly thicker than other v3-setae of similar length (Lienhard & Ferreira, 2014: fig. 
INNE) e REI, IT PT RN rina Psyllipsocus subtilis 
Forewing with a dark brown drop-shaped spot or stripe or a light brown diffuse patch in middle of cells rl and r3. 
Nfale Clin alrodSipresenMReMaAlE NS DIIOSVAUNLLONN ERP ne 29 
Forewing with a dark brown spot or stripe in middle of cells rl and r3. Male: phallic cradle of hypandrium 
anteriorly broadly rounded (Lienhard & Ferreira, 2014: figs 8, 9)... Psyllipsocus clunioventralis 
These cells only with a light brown diffuse patch covering most of their surface. Male: phallic cradle of hypandrium 
AMICnONhy tmumncalis (Liga nernel 6 Nearer, OISEAU) s000000500900056050000005000000000000000000000500030000000 Psyllipsocus didymus 


136 C. Lienhard & R. L. Ferreira 


GENERAL DISCUSSION 


Analysis of distribution 

This analysis is based on the data presented in Appendix 
2. Detailed considerations concerning each of the 19 
endemics (15 Psyllipsocus spp., 4 Neotrogla spp.) have 
already been published by Lienhard ef al. (2010b) and 
Lienhard & Ferreira (2013a, b, 2014) when describing 
these previously unknown species. During this 
project, about 400 adult individuals of 23 species of 
Psyllipsocidae and Prionoglarididae from 124 Brazilian 
caves were identified (Table 1). The total number 
of caves wherein each species was recorded and the 
number of caves wherein it was associated with another 
species of these families are presented in Table 2. The 
proportion of Brazilian caves occupied by these families 
is unknown; because of uneven sampling effort their 
absence cannot be assumed for caves where they are not 
currently recorded. This discussion only concerns caves 
with identified adults; caves with nymphs that cannot be 
identified are not included (see also Introduction). 

As previously observed by Lienhard & Ferreira (2014) and 
Yoshizawa et al. (2014), these insects live predominantly 
in dry resource-poor caves. Two closely related species 
of Psyllipsocus or two species of Neotrogla have never 
been found in the same cave. In most of the caves only 
one species was present, rarely two species (23 caves), 
exceptionally three species (viz. Gruta de Ubajara, CE: 
Psoc. lachlani, P. spinifer, P. yucatan; Gruta Janeläo, MG: 
N. brasiliensis, P. serrifer, P. spinifer; Gruta do Lagedo 
Grande, RN: P clunjunctus, P. spinifer, P. subtilis). In 
most caves with two or three species, only one of them is 
a Brazilian endemic; in seven caves two endemics were 
found and in two caves three endemics live together 
(Gruta Janelào and Gruta do Lagedo Grande, see above). 
In three caves (cave PEA 343, BA; Gruta Caboclo, MG; 
Gruta Janelao, MG) specimens of both genera, Neotrogla 
and Psyllipsocus, were present simultaneously. Among 
the 15 endemics of Psyllipsocus, 8 species were recorded 
from a single cave each (Table 2). Thus, very strict local 
endemism appears to be high in this genus. 

The cosmopolitan and often domicole species 
P. ramburii is the most common cave Psyllipsocus in 
Brazil (present in 30 caves, in 23 of them exclusively). 
In about 40% of the caves at least one of the non- 


endemics P. ramburii, P. yucatan, Psoc. lachlani or Psoc. 
pilipennis is present; 32% of the caves are exclusively 
inhabited by non-endemics. The high frequency of the 
opportunistic, parthenogenetic (thelytokous) and often 
domicole P ramburii indicates that this species is a 
particularly good disperser; but it may sometimes have 
been introduced by human activities. 

In all 9 caves with two or three Brazilian endemics at 
least one of them is P spinifer or P. clunjunctus. Together 
with P. serrifer, these are the most widely distributed 
endemics. P spinifer is known from 20 caves (19 
municipalities, 8 states), P clunjunctus from 16 caves (9 
municipalities, 5 states) and P. serrifer from 13 caves (9 
municipalities, 2 states) (for the distribution of the latter 
two species, see also map in Lienhard & Ferreira, 2013b). 
P. spinifer is very often associated with other species, 
being found alone in only five caves. It is interesting to 
see that the non-domicole, non-parthenogenetic endemic 
P. spinifer, the second most common Brazilian cave 
species, is much more often associated with a probably 
more specialized local endemic than the most common, 
cosmopolitan, domicole and parthenogenetic P ramburii 
(Table 2). It cannot be excluded that the presence of this 
opportunistic generalist contributes to the elimination 
of locally endemic specialists. Only in two of 30 caves 
P. ramburii has been recorded together with a Brazilian 
endemic (P falcifer, P. serrifer). 


Some evolutionary considerations 


This rather speculative discussion follows the short 
remarks that we made in the recent paper wherein most of 
the endemic cave species of Psyllipsocus were described 
(Lienhard & Ferreira, 2014). Rather than as a discussion 
of concrete results we consider this part as an outlook 
on possible future research perspectives concerning the 
fascinating cave psocids of the families Psyllipsocidae 
and Prionoglarididae. 

The only clearly troglobitic representative of these 
families from the New World is the prionoglaridid 
Speleopsocus chimanta known from a humid cave in 
south eastern Venezuela (see Key and Appendix 1). 
All other species are winged, have well developed 
compound eyes and are relatively well-pigmented, 
except for the cosmopolitan and often domicole pale and 


Table 1. Species richness, rate of endemism and distribution for Brazilian genera of Psyllipsocidae and Prionoglarididae based on adult 


specimens collected in caves. 


species endemics 
Psocathropos 2 
Psyllipsocus 17 15 (88%) 
Neotrogla 4 4 (100%) 


Total 23 19 (83%) 


caves municipalities states 
13 12 8 
102 51 13 
19 9 3 
124 59 13 


1577 


Brazilian cave psocids 


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138 C. Lienhard & R. L. Ferreira 


wingless forms of Psyllipsocus ramburii, with reduced 
but still multifacetted eyes (see Key). The only character 
interpreted as a cave adaptation (Lienhard, 2000) is 
the presence of leg trichobothria in the prionoglaridid 
subfamiliy Speleketorinae, to which the Brazilian genus 
Neotrogla belongs (Lienhard, 2004a; Lienhard er al, 
2010b). This genus is only known from caves. However, 
most of the Psyllipsocus species known from North and 
Middle America live on the bark of trees, in soil litter 
or on lichen-covered rock outcrops (Mockford, 2011). 
Unfortunately, all available data on Brazilian Psy/lipsocus 
concern material collected in caves, though some species 
of this genus might also live in other microhabitats in 
Brazil, outside of caves, as in the northern parts of the 
New World. It would be of particular interest to discover 
non-cavernicolous populations of the endemic species or 
of species closely related to them. 

Most of the local endemics are only known from one 
cave or from some caves situated close together (often in 
the same municipality or neighbouring municipalities). 
Some isolated relict populations of formerly more 
widely distributed species might be at the origin of this 
high diversity of cave endemics. These ancestors were 
probably not strictly cavernicolous. The relatively wide 
distribution of P spinifer, P. clunjunctus and P. serrifer 
may be the result of secondary dispersal of these 
endemics out of their region of origin. 

The endemic cave Psyllipsocus form a heterogeneous 
assemblage of isolated species or small groups of 
species. However, except for the most common endemic 
P. spinifer, they all show some kind of reduction of the 
phallosome, apparently due to homoplasy (Lienhard 
& Ferreira, 2014) (Note: males of P thaidis and 
P. radiopictus are not known). All known Brazilian 
prionoglaridids (1.e. the four species of the endemic cave 
genus Neotrogla) show also a strong reduction of the 
phallosome (Lienhard er al., 2010b; Lienhard & Ferreira, 
2013a; Yoshizawa et al., 2014). Therefore we proposed 
the speculative hypothesis that the locally endemic 
cave species of these genera might have evolved under 
a particular selection pressure favouring reduction of 
male primary external genitalia (Lienhard & Ferreira, 
2014). This selection pressure could be similar to that 
favouring reduction of the phallosome in the African 
cave prionoglaridid Afrotrogla, known from caves 
in semidesertic regions of Namibia and South Africa 
(Lienhard, 2007). All these psocids live in dry resource- 
poor caves and are the only known representatives of the 
suborder Trogiomorpha with reduced phallosomes. For 
the genus Neotrogla, Yoshizawa et al. (2014) showed 
that this reduction of the male intromittent organ is 
very probably related to reversed sexual selection. The 
only psocid species where reversed sexual selection has 
clearly been documented at present, Lepinotus patruelis, 
also belongs to the suborder Trogiomorpha but is not 
cavernicolous and has normal male genitalia (Wearing- 
Wilde, 1995, 1996; Yoshizawa et al., 2014). It would be 


interesting to elucidate the influence of a subterranean 
mode of life in dry caves on sexual selection. Miller & 
Svensson (2014: p. 427) “suggest that deeper ecological 
perspectives on sexual selection may alter some of the 
fundamental assumptions of sexual selection theory and 
rapidly lead to new discoveries”. Prionoglarididae and 
Psyllipsocidae could play a key role in future research 
in this field. We suggest that adaptations to life in dry, 
resource-poor cave environments could affect sexual 
selection and thus promote sexual role reversal in these 
insects. 

However, the absence of data on non-cavernicolous 
representatives of these families from Brazil makes it 
impossible to estimate the pertinence of the supposed 
correlation between a subterranean mode of life and the 
presence of reduced male genitalia. The most widely 
distributed endemic cave species is P. spinifer, the only 
endemic with a normal male phallosome, like that of 
all non-cavernicolous species of the genus. Thus, this 
species may also live outside of caves in Brazil. For 
the local endemics with reduced phallosome it may 
be speculated that they evolved as cave refugees from 
populations which were not competitive under out-of- 
cave conditions. 

In the trogiomorphan species with the most striking 
reduction of the phallosome (i.e. all species of Neotro- 
gla and Afrotrogla and of the Psyllipsocus clunjunctus 
species group), females have evolved novel differentia- 
tions of the spermapore region. Yoshizawa et al. (2014) 
showed that, in Neotrogla, the novel female organ func- 
tions as an intromittent organ during copulation. Such a 
function was also suggested hypothetically for the novel 
female organs in Afrotrogla (Lienhard, 2007) and the 
Psyllipsocus clunjunctus group (Lienhard & Ferreira, 
2013b). Among the Brazilian endemics with a reduced 
phallosome there are two particularly successful spe- 
cies, in terms of their distribution range, P clunjunctus 
and P. serrifer, both belonging to the clunjunctus spe- 
cies group. They are not only characterized by the almost 
complete absence of phallosome sclerites and the pre- 
sence of a novel female “micropenis”, but they also have 
evolved a novel male accessory genital organ, the clunial 
bridge (Lienhard & Ferreira, 2013b). Might these novel 
structures be the reason of their success? 

It is difficult to ascribe to pure coincidence these varied 
phenomena relating to similar modifications of genital 
morphology and reproductive behaviour in species that 
are not particularly closely related but which share a 
cavernicole life history. Perhaps we are here confronted 
with a new category of adaptations to a life in caves 
different to the classical troglomorphic adaptations 
(reduction of wings, pigmentation and eyes; see Dethier 
& Hubart, 2005), and as yet, not directly explained. 


Brazilian cave psocids 139 


ACKNOWLEDGEMENTS 


We thank Edward L. Mockford (Illinois State University, 
Normal, Illinois, USA) and John Hollier (MHNG) for 
critical reading of the manuscript and useful comments. 
CL thanks John Hollier for interesting discussions, 
Eliane De Coninck (Tervuren) for the loan of the type- 
series of Psyllipsocus collarti and Manuela Lienhard 
for composing the photo plate of Fig. 1. RLF is very 
grateful to all members of the team from the Centro 
de Estudos em Biologia Subterranea (UFLA) for their 
help in collecting material. Funding was provided by 
the Conselho Nacional de Pesquisa (CNPq - process n° 
477712/2006-1, and CNPq grant nr. 301061/2011-4) 
and the Fundaçäo de Amparo a Pesquisa do Estado de 
Minas Gerais (FAPEMIG - processes CRA - APQ 01826- 
08 and CRA - PPM-00433-11). 


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APPENDIX 1: Checklist of South American Psyllipsocidae 
and Prionoglarididae, with indication of general distribution 
(only known from caves, except for Psocathropos spp., 
Psyllipsocus delamarei, P. ramburii and P yucatan). 


Psyllipsocidae 

Psocathropos Ribaga, 1899: 156; type species: P lachlani 
Ribaga. 

Psocathropos lachlani Ribaga, 1899: 157. Widely distributed in 
the tropics (often domicole). 

Psocathropos pilipennis (Enderlein, 1931: 221). Widely 
distributed in the tropics (sometimes domicole). 


Psyllipsocus Selys-Longchamps, 1872: 145; type species: 
P ramburii Selys-Longchamps. 

Psyllipsocus angustipennis Lienhard, 2014; in: Lienhard & 
Ferreira, 2014: 239. Brazil. 

Psyllipsocus clunioventralis Lienhard, 2014; in: Lienhard & 
Ferreira, 2014: 225. Brazil. 

Psyllipsocus clunjunctus Lienhard, 2013; in: Lienhard & 
Ferreira, 2013b: 424. Brazil. 

Psyllipsocus delamarei Badonnel, 1962: 187. Argentina. 

Psyllipsocus didymus Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 229. Brazil. 

Psyllipsocus falcifer Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 216. Brazil. 

Psyllipsocus fuscistigma Lienhard, 2014; in: Lienhard & 
Ferreira, 2014: 233. Brazil. 

Psyllipsocus marconii Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 220. Brazil. 

Psyllipsocus proximus Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 242. Brazil. 

Psyllipsocus punctulatus Lienhard, 2014; in: Lienhard & 
Ferreira, 2014: 236. Brazil. 

Psyllipsocus radiopictus Lienhard, 2014; in: Lienhard & 
Ferreira, 2014: 234. Brazil. 

Psyllipsocus ramburii Selys-Longchamps, 1872: 146. 
Cosmopolitan (often domicole). 

Psyllipsocus serrifer Lienhard, 2013; in: Lienhard & Ferreira, 
2013b: 429. Brazil. 

Psyllipsocus similis Lienhard, 2013; in: Lienhard & Ferreira, 
2013b: 433. Brazil. 

Psyllipsocus spinifer Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 212. Brazil. 

Psyllipsocus subtilis Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 229. Brazil. 

Psyllipsocus thaidis Lienhard, 2014; in: Lienhard & Ferreira, 
2014: 223. Brazil. 

Psyllipsocus yucatan Gurney, 1943: 212. Widely distributed in 
the tropics. 


Prionoglarididae 

Neotrogla Lienhard, 2010; in: Lienhard ef al., 2010b: 612; type 
species: N. brasiliensis Lienhard. 

Neotrogla aurora Lienhard, 2010; in: Lienhard et al., 2010b: 
619. Brazil. 

Neotrogla brasiliensis Lienhard, 2010; in: Lienhard ef al., 
2010b: 614. Brazil. 

Neotrogla curvata Lienhard & Ferreira, 2013a: 4. Brazil. 

Neotrogla truncata Lienhard, 2010; in: Lienhard et al., 2010b: 
622. Brazil. 


Brazilian cave psocids 141 


Speleopsocus Lienhard, 2010; in: Lienhard ef al., 2010a: 186; 
type species: S. chimanta Lienhard. 

Speleopsocus chimanta Lienhard, 2010; in: Lienhard et al. 
2010a: 187. Venezuela. 


APPENDIX 2: Comprehensive list of Brazilian caves wherein 
adults of Psyllipsocidae and Prionoglarididae were collected 
(leg. R. L. Ferreira unless other collector mentioned) 


P. = Psyllipsocus; Psoc. = Psocathropos; in bold type = 
non-endemic species (widely distributed in the tropics or 
cosmopolitan) 


Alagoas (AL) 

Delmiro Gouveia (AL), Gruta Morcego, 23.iv.2006: Psoc. 
lachlani 

Murici (AL), Toca da Raposa 1 (granite), 13.1.2007: P 
radiopictus, P yucatan 

Murici (AL), Toca da Raposa 2 (granite), 13.1.2007: P yucatan 


Amazonas (AM) 
Apui (AM), Gruta Apiaca 1, Parque Nacional do Juruena, 
13.1x.2011: P proximus 


Bahia (BA) 

Campo Formoso (BA), Toca do Angico, 9.1.2008: P spinifer 

Campo Formoso (BA), Toca do Morrinho, i.1997: P spinifer 

Campo Formoso (BA), Toca do Pitu, 10.vii.2008: Neotrogla 
truncata 

Campo Formoso (BA), Toca da Tiquara, 8.1.2008, i.2009, 
x11.2010: P yucatan 

Curaçä (Patamuté) (BA), Toca d’agua de Patamuté, 6.1.2008: 
Psoc. lachlani, P. spinifer 

Ourolandia (BA), Toca dos Ossos, 31.vii.2007, 10.vi.2012, 
1.2013: Neotrogla truncata 

Palmeiras (BA), Gruta Ioiò, 1.1.2008: Neotrogla truncata 

Santa Maria da Vitoria (BA), cave PEA 341, x.2012: Neotrogla 
curvata 

Santa Maria da Vitoria (BA), cave PEA 342, x.2012: Neotrogla 
curvata 

Santa Maria da Vitoria (BA), cave PEA 343, 15.v.2011, leg. S. 
S. Salgado: Neotrogla curvata, P yucatan 

Sao Desidério (BA), Gruta do Catitu, 24.vii.200: P clunjunctus 

Sao Desidério (BA), Gruta do Sumidouro do Joao Baio, 
29.vii.2006: P clunjunctus, P. spinifer 

Sao Félix do Coribe (BA), cave PEA 377, 10.v.2011, leg. S. S. 
Salgado: P serrifer 

Sao Félix do Coribe (BA), cave PEA 378, x.2012: Neotrogla 
curvata 

Sao Félix do Coribe (BA), cave PEA 380, 21.vii.2011, leg. S.S. 
Salgado: Neotrogla curvata 

Sao Félix do Coribe (BA), cave PEA 381, 18.vii.2011, leg. S.S. 
Salgado: Neotrogla curvata 

Sao Félix do Coribe (BA), cave PEA 383, 18.vii.2011, leg. S. S. 
Salgado: Neotrogla curvata 

Varzea Nova (BA), Gruta Jurema, 20.vii.2008: P clunjunctus 


Ceara (CE) 

Araripe (CE), Gruta do Brejinho, 1.v.2007: P spinifer 

Tejuçuoca (CE), Gruta do Veado Campeiro, 16.ix.2008: P 
fuscistigma, P. spinifer 


Ubajara (CE), Gruta do Araticum, 1.1.2007: Psoc. pilipennis, 
P. spinifer 

Ubajara (CE), Gruta dos Mocés, 3.1.2007: P yucatan 

Ubajara (CE), Gruta do Morcego Branco, 3.1.2007: P spinifer, 
P yucatan 

Ubajara (CE), Gruta de Ubajara, 30.xii.2006: Psoc. lachlani, P. 
spinifer, P yucatan 


Espirito Santo (ES) 

Castelo (ES), Gruta do Limoeiro, 7.1.2005: P ramburii, P 
yucatan 

Santa Teresa (ES), Gruta do André Huscki, 4.1.2005: Psoc. 
lachlani 


Goias (GO) 
Damianopolis (GO), Lapa do Ribeirao dos Porcos, 29.vii.2001, 
5.x.2001, 27.vi.2002: P clunjunctus, P. spinifer 


Minas Gerais (MG) 

Arcos (MG), Caverna do Alinhamento, 1.vi.2002: P ramburii 

Arcos (MG), Gruta da Bocaininha, 3.x11.2008: P serrifer 

Arcos (MG), Gruta Labirinto, 28.1.2006: P serrifer 

Arinos (MG), Lapa do Salobo, 18.vii.2010: P yucatan 

Cordisburgo (MG), Gruta de Maquiné, 10.vii.2000, 18.v.2010: 
Psoc. lachlani, P ramburii 

Cordisburgo (MG), Gruta do Salitre, 22.iv.2011: P serrifer 

Cordisburgo (MG), Gruta Santo Amaro 1, 28.1x.2010: P 
ramburii 

Cordisburgo (MG), Gruta Tao Lucas, 14.x1.2010: P falcifer 

Cordisburgo (MG), Lapinha do Atamis, 13.xi.2010: P falcifer 

Coromandel (MG), Gruta Joao do P6, 5.x.2000: P ramburii 

Coromandel (MG), Gruta Ronan, 3.x.2000: P ramburii 

Coromandel (MG), Gruta Ronan II, 4.x.2000: P ramburii 

Diamantina (MG), Gruta do Salitre, 13.x11.2007: P ramburii 

Doresopolis (MG), Gruta P43, 9.xi.2003: P serrifer 

Itabirito (MG), Gruta MP1, 29.viii.2005: P serrifer 

Itabirito (MG), Gruta MP8, 8.1x.2005: P similis 

Itacarambi (MG), Gruta Bonita, 19.111.2003: P angustipennis 

Itambé do Mato Dentro (MG), Baixada dos Crioulos 2, 
29.vii.2004: P similis 

Itumirim (MG), Gruta Santo Antonio, 25.x.2002: P ramburii 

Januaria (MG), Gruta Caboclo, 27.vii.2003: Neotrogla 
brasiliensis, P. spinifer 

Januäria (MG), Gruta Ossos, vii.2003, iii.2013: Neotrogla 
brasiliensis 

Januaria / Itacarambi (MG), Gruta Brejal, 25.vii.2003: female 
cf. P punctulatus 

Januäria / Itacarambi (MG), Gruta Janeläo, 28.vii.2003: 
Neotrogla brasiliensis, P serrifer, P spinifer 

Januäria / Itacarambi (MG), Gruta Preguiça, 26.vii.2003: P 
angustipennis 

Joao Pinheiro (MG), Gruta do Sapecado, 15.x.2010: P ramburii 

Lagoa da Prata (MG), Gruta Saläo de Festas, 4.v.2003: P 
serrifer 

Lagoa Santa (MG), Gruta da Lapinha, 12.vi.2002: P ramburii 

Matozinhos (MG), Gruta dos Irmäos Piriäs, 28.vii.2000: P 

ramburii 

Matozinhos (MG), Gruta Lavoura, 28.+ 29.v.1997, 26.vi.1997, 

19.11.2000: P ramburii 

Matozinhos (MG), Gruta Pequenas III, 31.vii.2002: P serrifer 

Matozinhos (MG), Meandro Abismante, 6.vii.2002: P ramburii 

Matutina (MG), Gruta 9, 10.x.2010: PR ramburii 

Moeda (MG), SMS 19, 3.xii.2005: P similis 


142 C. Lienhard & R. L. Ferreira 


Moeda (MG), SMS 29, 11.xii.2005: P ramburii 

Montalvania (MG), Gruta Nossa Senhora do Perpétuo Socorro, 
14.vii.2007: Psoc. lachlani, P marconii 

Nacip Raydan (MG), Gruta Manga de Pedra (granito), 
21.vii.2002: Psoc. lachlani 

Pains (MG), Buraco do Nando, 12.x.2003: P serrifer 

Pains (MG), Gruta Brasical, 28.ix.2003: P serrifer 

Pains (MG), Gruta do Capào, 5.v.2001: P ramburii 

Pains (MG), Gruta dos Estromatolitos, 7.xi.2000: P falcifer 

Pains (MG), Gruta Paiol de Milho, 13.x.2003: P falcifer, P. 

spinifer 

Pains (MG), Gruta Paranoa, 15.1.2008: P ramburii, P. serrifer 

Pains (MG), Gruta Retiro, 27.xi.1999: P ramburii 

Pains (MG), Gruta Ronco, 28.xi.1999: P falcifer 

Pains (MG), Gruta do Sobradinho, 5.v.2001: P. serrifer 

Paracatu (MG), Gruta da Fazenda Tamandua II, 14.x.2010: P 
ramburii 

Paracatu (MG), Lapa do Brocotò, 16.ix.2010: P ramburii 

Paracatu (MG), Lapa de Santo Antonio, 13.vii.2010: P 
ramburii 

Presidente Olegario (MG), Lapa Vereda da Palha, 13.x.2010: 
P. clunjunctus 

Santa Maria do Suaçui (MG), Gruta do Rio Suacui, 19.vii.2002: 
P ramburii, P yucatan 

Sete Lagoas (MG), Gruta Rei do Mato, 3.+4.xi.2011: P falcifer, 
P ramburii 

Teofilo Otoni (MG), Lapa da Vaca Parida, 26.1.2005: Psoc. 
lachlani, P. yucatan 

Vazante (MG), Gruta da Escarpa, x1.2008: P falcifer 

Vazante (MG), Lapa da Delza, 12.vii.2010: P ramburii 

Vazante (MG), Lapa das Urtigas, 16.ix.2010: P falcifer 


Mato Grosso (MT) 

Apiacas (MT), Casa de pedra do Navalha, Parque Nacional do 
Juruena, 9.ix.2011: P angustipennis 

Chapada dos Guimaraes (MT), Gruta Kiogo Brado, 27.x.2006: 
P. clunioventralis, P. spinifer 

Paranaita (MT), Gruta da Pedra Preta, 19.ix.2011: P didymus 


Piaui (PI) 

Coronel José Dias (PI), Coroa de Frade, 1x.2008: P thaidis 

Coronel José Dias (PI), Toca do Garrincho, 11.ix.2008: Psoc. 
pilipennis 

Coronel José Dias (PI), Toca do Inferno, 12.ix.2008: P 
punctulatus, P. spinifer 

Coronel José Dias (PI), Toca das Moendas, 10.ix.2008: P 
spinifer 


Rio Grande do Norte (RN) 

Baraùna (RN), Caverna Britador, 11.vi.2010, leg. D. M. Bento: 
P. clunjunctus 

Barauna (RN), Caverna Cipös, 11.vi.2010, leg. D. M. Bento: 
P clunjunctus 

Baraüna (RN), Caverna Escada, 27.1.2010, leg. D. M. Bento: 
P. clunjunctus 

Baraüna (RN), Caverna Esquecida, 17.vi.2010, leg. D. M. 
Bento: P clunjunctus 


Felipe Guerra (RN), Caverna Arapua, 3.viii.2010, leg. D. M. 
Bento: P subtilis 

Felipe Guerra (RN), Caverna Beira-Rio, 19.11.2010, leg. D. M. 
Bento: P subtilis 

Felipe Guerra (RN), Caverna Rumana, 19.1.2010, 5.viii.2010, 
leg. D. M. Bento: P clunjunctus, P subtilis 

Felipe Guerra (RN), Caverna Trapia, 6.1.2010, 4.viii.2010, leg. 
D. M. Bento: P yucatan 

Felipe Guerra (RN), Gruta Carrapateira, 24.iv.2007: P 


clunjunctus 

Felipe Guerra (RN), Gruta da Catedral, 14.ix.2008: P 
clunjunctus 

Felipe Guerra (RN), Gruta Roncador, 1.vi.2006: Psoc. 
pilipennis 


Felipe Guerra (RN), Lapa do Engano, 5.viii.2010, leg. D. M. 
Bento: P clunjunctus 

Governador Dix-Sept Rosado (RN), Caverna Capoeira do Joao 
Carlos, 3.vi.2010, leg. D. M. Bento: P subtilis 

Governador Dix-Sept Rosado (RN), Gruta do Lagedo Grande, 
21.vii.2010, leg. D. M. Bento: P clunjunctus, P. spinifer. 
P subtilis 

Governador Dix-Sept Rosado (RN), Gruta do Marimbondo 
Caboclo, 20.vii.2010, leg. D. M. Bento: P subtilis 

Jandaira (RN), Gruta Aroeira, 11.1.2006: Psoc. pilipennis 

Mossorò (RN), Caverna Trinta, 10.vi.2010, leg. D. M. Bento: 
P. clunjunctus 


Rio Grande do Sul (RS) 
Torres (RS), Furna da Lagoa de Itapeva, 20.v.2008: P ramburii 


Sao Paulo (SP) 

Altinépolis (SP), Gruta Edgar 1, 28.11.2006: P spinifer 

Altinépolis (SP), Gruta do Itambé, 27.11.2006: Psoc. lachlani, 
P ramburii 

Altinöpolis (SP), Gruta Olho de Cabra, 2.111.2006: P ramburii, 
P spinifer 

Altinépolis (SP), Gruta do Parana, 1.111.2006: P ramburii, P. 
spinifer 

Ilha Bela (SP), Gruta da Serraria, 22.iv.2006: P yucatan 

Itirapina (SP), Gruta da Toca, 22.x.2004: P ramburii 


Tocantins (TO) 

Arraias (TO), cave Ponto 008, 4.-6.x.2010, leg F. Pellegatti ef 
al.: Neotrogla aurora 

Arraias (TO), cave Ponto 014, 4.-6.x.2010, leg F. Pellegatti er 
al.: Neotrogla aurora 

Aurora do Tocantins (TO), Gruta Asa Branca 1, 7.i.2009, leg. 
R. A. Zampaulo: Neotrogla aurora 

Aurora do Tocantins (TO), Gruta Biritite, 5.1.2009, leg. R. A. 
Zampaulo: Neotrogla aurora 

Aurora do Tocantins (TO), Gruta Couve-Flor, 7.1.2009, leg. R. 
A. Zampaulo: Neotrogla aurora 

Aurora do Tocantins (TO), Gruta das Ras, 8.1.2009, leg. R. A. 
Zampaulo: P. clunjunctus 

Dianöpolis (TO), Gruta Imperial, iii.2013: Neotrogla aurora 


Revue suisse de Zoologie (March 2015) 122(1): 143-148 ISSN 0035-418 


Live trapping design for the harvest mouse (Micromys minutus) in its summer habitat 


Peter Vogel!‘ & Antoine Gander? 


! Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland. 
Association de la Grande Cariçaie, Chemin de la Carigaie 3, 1400 Cheseau-Noreaz, Switzerland. 
E-mail: a.gander@grande-caricaie.ch 


Abstract: The harvest mouse Micromys minutus is a very rare species in Switzerland. It has only been documented 
accurately since 1960. Most records are based on nest findings and there have been few direct observations or captures, 
mainly because live trapping of this species is not simple. Therefore, an efficient trapping technique is needed for 
population studies and to facilitate the management of its habitat. By combining the methods used to capture very small 
(Suncus etruscus) and climbing (Muscardinus avellanarius) mammals, we developed a design using Longworth traps 
with mouse excluders set on suspended platforms. This allowed us to trap more harvest mice in four field sessions of 60 
trap-nights than have ever been caught previously since its discovery in Switzerland. 


Keywords: Harvest mouse, Longworth trap, mouse excluder, prebaiting. 


Resumé: La souris des moissons (Micromys minutus) est une espèce très rare en Suisse et peu documentée jusque 
dans les années 1960. La plupart des indications de présence sont indirectes, basées sur la découverte de nids. Très peu 
d’entre-elles font référence à des observations directes, qu’ elles soient visuelles ou issues de captures d’individus vivants, 
car le piégeage classique n’est pas efficace. La vérification de la bonne gestion de son habitat ou la réalisation d’études 
populationnelles nécessitent cependant des techniques de piégeage efficientes. Quelques astuces développées pour piéger 
de petites musaraignes (Suncus etruscus) et des muscardins (Muscardinus avellanarius) exploitant les structures hautes 
de la végétation ont aidé a développer un protocole ayant permis de piéger en quatre sessions de 60 nuits-pieges, plus de 
souris de moissons que jamais depuis sa découverte en Suisse. 


Mots-clés : Souris des moissons, piège Longworth, réducteur de la taille de l’entrée, pré-appater. 


INTRODUCTION War (OFEFP, 1990) and hence only a few regions in this 
country harbour this species. Thirty years ago, a few 
small populations were recorded in the southern part of 
Switzerland (Lardelli, 1981), the north-western area close 
to Lake Constance, the region of Basel (contiguous with 
the population in Alsace, France), the Geneva region and 
the most important population along the southern shore 


of Lake Neuchatel (Rahm, 1995). 


The harvest mouse Micromys minutus (Pallas, 1771) is a 
very rare rodent in Switzerland (Fig. 1). In older literature, 
it was mentioned only twice from the region of St-Gall 
(Fatio, 1869; Miller, 1812). Baumann (1949) and Hainard 
(1949) had no knowledge of its occurrence in Switzerland. 
In western Switzerland, it was first documented by Krapp 
(1964). This rare status is attributable to the altitudinal 


and climatic conditions, as well as the scarce occurrence 
of swampy habitats. This species occurring from Europe 
to Japan in a homogenous genetic clade (Yasuda er al., 
2005) is well distributed in most neighbouring countries, 
such as France, Germany and northern Italy (humid 
plain of the Po), but it is absent from the Alps, including 
many parts of Austria (Spitzenberger, 1986). The main 
habitat of M. minutus comprises reed beds in wetlands 
(Spitzenberger, 1999), approximately 90% of which have 
been destroyed in Switzerland since the Second World 


Manuscript accepted 6.10.2014 
* Deceased January 2015 
DOI: 10.5281/zenodo.14574 


Most records are based on indirect signs, i.e., the 
presence of its summer nests (Piechocki, 1958) woven 
with longitudinally spliced leaves (JuSkaitis & Remeisis, 
2007) within the dense vegetation of Cyperaceae (e.g., 
Carex spp.) or Poaceae (e.g. Phragmites communis and 
Phalaris arundinacea) species. Recent methodological 
developments of nest search (Blant er al., 2012) were 
applied to potential habitats, which detected new 
populations in the Ajoie region (Canton of Jura, north- 
western Switzerland), and extinctions were suggested 


144 P. Vogel & A. Gander 


Fig. 1. Harvest mouse captured on 7 March 2014 (Photo: P. Vogel). 


in many previously occupied localities where no recent 
observations could be obtained, e.g. southern Switzerland 
(Maddalena & Zanini, 2008) and Geneva (Blant ei al., 
2012). 

The more direct technique of trapping has not been used 
often because this species is rather difficult to catch 
with standard trapping designs, as mentioned in several 
reports (e.g., Piechocki, 1958; Trout, 1978; Rahm, 1995; 
Serrano Padilla, 1998). In the city of Oxford, Dickman 
(1986) found 26 nests but trapped only four M. minutus 
during 3858 trap nights. Data were checked at the 
Swiss Fauna Database (CSCF-Infofauna, www.cscf. 
ch) and among 201 occurrences based on known census 
techniques (excluding our own data), only five were 
captured with small mammal traps. All other records 
were based on nest sightings (135), owl pellet analyses 
(25), direct observations, mummified remains and foot 
prints (seven). Moreover, between 1996 and 2002, 27 
individuals were collected at Lake Neuchatel, which 


had drowned in the plastic buckets combined with drift 
fences used for regular amphibian censuses, a regrettable 
case of collateral damage for this rare species. Thus, 
in order to increase the success of live trapping, we 
developed a method that allowed us to capture 48 harvest 
mice and the present study reports the final standard trap 
design, which was applied to a small study of habitat 
management for this species (Vogel & Gander, in press). 
The method was inspired by trapping techniques 
developed for the small Etruscan shrew Suncus etruscus 
(Vogel, 2012) using long prebaiting periods with an 
entrance filter, and for the arboreal hazel dormouse 
Muscardinus avellanarius (Vogel et al., 2012) using 
hanging platforms to set the traps. Live trapping may 
facilitate different aspects of investigation such as 
density, home range and optimal habitat studies, which 
are important for the conservation of this rare species. 


Life trapping design for the harvest mouse 145 


MATERIAL AND METHODS 


Trapping locality: A trapping census using different 
trapping methods was performed at three localities on 
the southern shore of Lake Neuchâtel where M. minutus 
nests have been found (pers. obs.): Cheyres, Font 
and Portalban. This allowed us to find an important 
population in Font (lat. 46.83674°, long. 6.810381°, at 
430 m a.s.l.) at the end of the summer in 2012. Therefore, 
we restricted our further investigations to this locality, 
where we repeated the census in March 2013, September/ 
October 2013 and February/March 2014. 


Habitat: Three trap lines were set in the following 
habitats. Line | was in a habitat with a mix of Phalaris 
arundinacea and Phragmites communis. Line 2 was in 
an adjacent field of P communis in the direction of the 
lake between an Alnus glutinosa stand and a dune close 
to the water. Line 3 was located at a distance of 300 m 
in a mixture of Cladium mariscus and Carex panicea 
with some P communis standing in part in the water. 
During the first winter session, only line 2 remained in 
exactly the same place. As the reeds on line | and parts 
of line 3 were mowed during December 2012 for habitat 
conservation, line 1b was set in the adjacent forest and 


line 3b was shifted to the closest unmown area during 
March 2013. 


Trap design: For each line, we used 20 Longworth 
traps (Penlon Ltd, Abingdon, UK). They were set in 
alternating pairs, where one pair was placed on the 
ground and the next was placed at a height of about 
80 cm in the vegetation (Fig. 2). We used a wooden 
platform (13 x 33 cm) to fix the hanging traps within the 
vegetation, which was usually attached to Phragmites or 
young A/nus to ensure easy access by mice. One trap of 
each pair (odd numbers) had a normal entrance whereas 
the second trap (even numbers) had a reduced entrance 
by use of a mouse excluder (Fig. 3). This accessory 
equipment (Penlon) originally contained a circular hole 
of 12 mm and it was designed for increasing trapping 
success of shrews by avoiding trapping larger and 
far more common mice and voles. In our study, we 
increased the diameter of mouse excluders to 14 mm. 
Moreover, many holes were increased in size by 
gnawing mice. 


During the winter of 2013, the trap success on the 
platforms was very low demonstrating a drastic 
reduction of the climbing behaviour. Therefore, during 


> 
a 
Lo 


EYES SN 


ru Tai vai a 


2 
l 


rile 
ZEN) 
\i 


SIA Il 


Fig. 2. Top line: hanging platforms in summer and in winter; bottom line: traps on the ground in autumn and winter (Photos: P. Vogel). 


146 P. Vogel & A. Gander 


Fig. 3. Harvest mouse looking through the hole of a mouse excluder in a Longworth trap (Photo: P. Vogel). 


the winter of 2014, all 10 pairs of traps in each line were 
placed on the ground. 


The traps were prebaited for 2-4 nights (avoiding heavy 
rainfall) and then triggered for one night, with two or 
three checks per night. The bait was a mixture of seeds 
(sunflower, -wheat and millet) and mealworms, but a 
small piece of apple was included during the trapping 
nights. Initially, some seeds were scattered around the 
entrances to increase interest in the traps. 


The small mammals were not marked to avoid 
damaging M. minutus. Therefore, the number of 
controlled individuals of all species may include some 
recaptures. As no invasive method was used, we could 
not always distinguish the syntopic sibling species, 
Sorex araneus and Sorex coronatus. 


Statistics: We used chi-squared tests (X?) to compare 
the separate effects of the entrance diameter size on 
the trapping of shrews, harvest mice and other rodents 
combined, but only significant differences are reported. 
The same test was used for the two trap positions. We 
also separated the data according to the seasons. 


RESULTS AND DISCUSSION 


In total, 184 small mammals were captured during the 
four trapping sessions (Table 1), where 60 traps were 
each triggered for one night after prebaiting, which 
corresponded to 240 trap nights. The trap success rate 
(77%) was higher than that reported in other studies of 
harvest mice, e.g. 14.3% in the total catch (50,500 trap 
nights) and 37.7% in a reduced set using a standard design 
by Trout (1976), 53% by Nordwig e? al. (2001) and 12% 
by Haberl & Krystufek (2003). Among the eight species 
captured, Myodes glareolus (56 captures) was dominant, 
followed by Apodemus sylvaticus (39 captures) and 
M. minutus (34 captures). For the seasonal score, in the 
summer 2012 trapping session, the dominant species 
were M. minutus together with M. glareolus (14 captures 
each). The harvest mouse disappeared during the winter 
and none were trapped in March 2013. However during 
October 2013, the habitat had a similar harvest mouse 
population density (number caught per trap line) as that 
recorded in the previous year and was the dominant 
species (19 captures). In the very mild winter of 2014, 
one harvest mouse was captured. 


Life trapping design for the harvest mouse 


147 


Table 1. Summary ofthe four trapping sessions (the three habitats combined), which shows the numbers of rodents and shrews trapped 
on the ground (G) and platforms (P), and the numbers with the normal (N) and reduced (R) entrances. During the winter of 


2014, all of the trap pairs were placed on the ground. This explains the twofold increase compared with the winter of 2013. 


Season Summer 2012 
Species/Trap type G30 P30 N30 R30 G30 
Sorex araneus/coron. 3 0 2 i 10 
Sorex. minutus 0 0 0 0 3 
Micromys minutus 3 11 8 6 0 
Apodemus flavicollis 2 4 6 0 0 
Apodemus sylvaticus 2 5 4 3 2 
Myodes. glareolus 8 6 6 8 8 
Microtus agrestis 3 l 2 2 3 
Total 21 27 28 20 26 


The comparison of trap preferences, 1.e., ground versus 
platform, showed that M. minutus was trapped more 
often in the platform traps than the ground traps during 
the summer of 2012 (11 versus three, respectively, 
but the difference was not significant), whereas the 
numbers were almost equal during the autumn of 2013 
(10 versus nine). This is probably because the climbing 
activity of M. minutus is reduced in the autumn when 
a drastic change in habitat exploitation occurs. Nordvig 
et al. (2001) tested traps on the ground and attached to 
vegetation in September and obtained similar results (13 
M. minutus captured in elevated traps but only three on 
the ground). It was concluded that the summer decline 
in captures by traps placed exclusively on the ground 
mentioned by Trout (1978) may have been a consequence 
of greater activity in elevated vegetation. 

In the three seasons with platforms, about 2/3 of the other 
small mammals were trapped more often on the ground 
than on the platforms during summer and autumn (32 
versus 24), although this difference was not significant. 
However, during the winter of 2013 (26 versus three), the 
traps on the ground captured significantly more (x? = 9.9, 
df= 1, P< 0.001). The sunflower seeds scattered around 
the traps on the platform were not touched and even birds 
did not visit them, except for one Parus palustris. 

In the trap entrance size comparison, i.e., normal (N) 
versus reduced (R), we expected significant differences 
between the three categories: 1) shrews, 2) M. minutus 
and 3) other larger rodents. However, shrews (15 N, 
20 R) and M. minutus (18 N, 16 R) did not exhibit 
significant preferences. It is even possible that normal 
(larger) entrances are preferred by M minutus, but 
these traps were frequently occupied by larger species. 
Indeed, the trap occupation rate by larger rodents was 
much higher in traps with normal entrances (84 N) than 
reduced entrances (31 R) and the difference was highly 
significant (4? = 11.75, df = 1, P < 0.001). Trout (1976) 
subdivided his study area into a grid of 10 x 10 m and 
placed four Longworth traps in the centre of each grid 


Winter 2013 Autumn 2013 Winter 2014 
P30 N30 R30 G30 P30 N30 R30 N30 R30 
0 5 5 6 0 4 2 2 10 

0 2 1 0 0 0 0 0 | 

0 0 0 9 10 9 10 1 0 

0 0 0 0 0 0 0 0 0 

0 ? 0 5 7 8 4 16 2 

2 6 4 3 ] 2 2 25 5 

l 3 l 0 0 0 0 4 0 

3 18 11 23 18 23 18 48 18 


cell, where two traps were normal and two contained a 
mouse excluder with a 13-mm hole. By recalculating 
his data, we found that significantly more M. minutus 
were captured in the traps with reduced entrance sizes 
(£ = 17.6, df= 1, P< 0.001). The trap type, trap density, 
small mammal species community and species density 
have each potential strong effects on the result. The 
effect of trap type was demonstrated by Serrano Padilla 
(1998) who used a large enclosure with an exclusively 
experimental harvest mouse population where Ugglan 
traps had a higher success (12.9%) than Longworth 
traps (2.7%), followed by Sherman traps (1.8%). The 
better capture rate of Ugglan traps may be explained 
by the rather “open” system and the possibility of 
multiple catches. In a natural environment however 
with a complex small mammal community Ugglan traps 
showed the same score for the harvest mouse compared 
to Longworth traps (PV pers. observation in Denmark). 
In conclusion, the use of double traps with reduced 
and normal entrance set on platforms appears to be the 
optimal trap design for M. minutus in the summer. Traps 
with reduced entrances were avoided by the majority 
of the more frequent larger rodents so the chance of 
capturing M. minutus in an unoccupied trap was higher. 
In addition, the by-catches provided an idea of the other 
species that shared the same habitat. 

The lack of M minutus captures during the winter 
suggests that there was a drastic change in their 
behaviour, either by subterranean habitat exploitation or 
a total habitat change. Previous studies of this issue lack 
agreement (Piechocki, 1958; Böhme, 1978). However, 
the use of a drift fence to sample amphibians resulted 
in high number of M. minutus captured, which suggests 
that migration to other habitats may occur (Koskela & 
Viro, 1976). However, in contrast to frogs, the direction 
of dispersion was not determined. This was also not the 
case for the 27 kills of harvest mice in amphibian pitfall 
traps from lake Neuchatel (in 27,293 traps nights, trap 
success 0.1%) mentioned in the introduction. Another 


148 P. Vogel & A. Gander 


possibility of lack of winter observation is a very high 
winter mortality where only a few individuals survive, 
but without any change of habitat use (Trout, 1978). To 
facilitate optimal habitat management, it would be useful 
to develop an adapted winter trapping technique. 


ACKNOWLEDGEMENTS 


We are very thankful to Simon Capt for extracting the 
harvest mouse data from the Centre Suisse de Cartogra- 
phie de la Faune, Neuchâtel, and for his comments on 
the manuscript. The Canton of Fribourg kindly provided 
the authorisation for trapping. Heinz Durrer, Tiziano 
Maddalena, André Meylan, Manuel Ruedi, Claude 
Vaucher, Darius Weber and Raffael Winkler helped to 
check problematic data of the list of historic occurrences 
of harvest mice in Switzerland. 

This study is dedicated to Urs Rahm (1925-2013) and 
Paul Marchesi (1958-2013), two enthusiastic Swiss 
mammalogists. 


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Revue suisse de Zoologie (March 2015) 122(1): 149-163 ISSN 0035-418 


Tapeworms (Cestoda: Proteocephalidea) of teleost fishes from the Amazon River in Peru: 
additional records as an evidence of unexplored species diversity 


Alain de Chambrier!, Roman Kuchta? & Tomas Scholz?” 


! Departement des Invertébrés, Muséum d'histoire naturelle, PO Box 6434, CH-1211 Geneva 6, Switzerland. 

E-mail: alain.dechambrier@ville-ge.ch 

Institute of Parasitology, Academy of Sciences of the Czech Republic, Branisovska 31, 370 05 Ceské Budejovice, 
Czech Republic. 

Corresponding author. E-mail: tscholz@paru.cas.cz 


Abstract: This paper represents an update of the previous list of adult proteocephalidean tapeworms (Cestoda) parasitizing 
freshwater teleosts from the Peruvian Amazon, which was presented by de Chambrier ef al. (2006a). Four new samplings 
made it possible to almost double the number of species found, all of them representing new geographical records 
from Peru. With 34 newly added species, a total of 63 proteocephalidean cestodes (46 named species of 27 genera) are 
now reported from Amazonia in Peru (compared to 54 named species of 28 genera from its Brazilian part). The genera 
previously unreported by de Chambrier et al. (2006a) are Ageneiella, Brayela, Endorchis, Ephedrocephalus, Gibsoniela, 
Harriscolex, Jauella, Lenhataenia, Manaosia, and Megathylacus. Four species, namely Jauella glandicephalus, 
Monticellia belavistensis, M. santafesina, and Proteocephalus hobergi, are reported from the Amazon River basin for 
the first time. Harriscolex piramutab (Woodland, 1934) n. comb. is proposed for specimens previously identified as 
Proteocephalus piramutab Woodland, 1934 from Brachyplatystoma vaillantii. The highest number of proteocephalidean 
cestodes is reported from Pseudoplatystoma fasciatum (a total of 10 cestode species), Zungaro zungaro (previously 
named Paulicea luetkeni; 9 species) and Phractocephalus hemioliopterus (6 species). A high number of unnamed species 
found in Peru (17), which most probably represent taxa new to science including at least two new genera, demonstrates 
that the species richness of proteocephalidean cestodes in Amazonia is still poorly known. 


Keywords: Catfish - freshwater fish - Siluriformes - Peru - Pimelodidae - Amazonia - species diversity - faunal survey. 


INTRODUCTION 


The Amazon River basin is by far the largest river basin 
in the world and drains roughly 40 percent of the South 
American continent. This river basin hosts the most diverse 
fish fauna in planet, with about 2,500 species described 
and another 1,000 species estimated to be described (Junk 
et al., 2007). However, recent large-scale environmental 
degradation due to anthropogenic pressure such as 
deforestation, water pollution, overfishing and intensive 
farming has had negative effect on water ecosystems, 
including considerable decrease of population density 
of big pimelodid catfishes (Siluriformes) in the Brazilian 
part of Amazonia (Angelini er al., 2006; Pelicice & 
Agostinho, 2008; Boni er al., 2011; Reis, 2013). 

These fishes serve as definitive hosts of a unique rich 
fauna of proteocephalidean tapeworms (Cestoda) that 
probably underwent explosive radiation and represent 
an interesting model for co-evolutionary studies because 
of strict host specificity of most taxa (de Chambrier & 
Vaucher, 1997, 1999; Zehnder & Mariaux, 1999; de 


Manuscript accepted 3.11.2014 
DOI: 10.5281/zenodo.14580 


Chambrier er al., 2004a; HypSa er al., 2005). Unlike most 
parts of the Brazilian Amazonia, in which population 
density of big catfish has declined (Angelini ef al., 2006; 
Pelicice & Agostinho, 2008), numerous stocks of these 
fishes including pimelodids still inhabit the Peruvian part 
of the Amazon River basins. 

Parasitological examination of 276 fishes of 73 species 
from the Amazon River and its tributaries around Iquitos, 
Loreto Region in Peru, carried out by the present authors 
and their co-workers in 2004 and 2005, revealed an 
extraordinary richness of proteocephalidean tapeworms, 
which were all reported from Peru for the first time (de 
Chambrier & Scholz, 2005; de Chambrier er al., 2006a). 
De Chambrier ef al. (2006a) listed as many as 29 species 
of 17 genera found in 10 species of siluriform fishes and 
cichlids. New sampling in this region in 2006, 2008, 2009 
and 2011 made it possible to obtain additional material 
that includes many cestodes not having been previously 
reported from Peru or from the Amazon River basin; some 
may even be new to science. To provide a robust baseline 
for forthcoming analyses of zoogeographical patterns 


150 A. de Chambrier, R. Kuchta & T. Scholz 


and phylogenetic relationships of proteocephalideans in 
the Neotropical Region, updated information is presented 
on the species composition, host-parasite associations 
and geographical distribution of these cestodes, which 
represent an important component of the parasite fauna 
of Neotropical fishes (Thatcher, 2006). 


MATERIAL AND METHODS 


A total of 611 fish from the Amazon River and its 
tributaries around Iquitos (72°50’-73°40’W; 3°34°- 
4°53’S), Loreto Region, Peru, were examined for 
parasites in September 2006 and 2008, and in October 
2009 and 2011. Intestines of freshly captured hosts 
as well as those sold on the market of Belén in Iquitos 
were transported in coolers to the provisional laboratory 
(courtesy of Acuario Rio Momön in Iquitos), where 
they were immediately examined. For morphological 
evaluation, only specimens in good condition were 
used, but quantitative parameters such as intensity or 
abundance could not be reliable assessed. 

Cestodes were gently washed in saline, fixed with 
hot (almost boiling) 4% formaldehyde solution and 
then processed using standard procedure used for 
fish tapeworms as described by de Chambrier er al. 
(2014). Fragments of strobila were also fixed with 96% 
molecular-grade ethanol for molecular analyses (DNA 
sequencing), which forms part of a large-scale study on 
the phylogenetic relationships of cestodes supported by 
the National Science Foundation project (programme 
Planetary Biodiversity Inventory; see www.tapeworm. 
uconn.edu). In the present paper, scanning electron 
micrographs (SEM) of the scoleces of five species are 
provided (Figs 1-5); these species were not studied using 
SEM or their SEM pictures were based on contracted or 
deformed specimens. 

Most specimens found are deposited in the Natural 
History Museum, Geneva, Switzerland (MHNG-PLAT), 
which hosts one of the most comprehensive collections 
of proteocephalidean cestodes (http://www.ville-ge.ch/ 
mhng/dpt inve coll e.php#platyhelminthes). See Table 
1 for more details. Classification of cestodes, including 
original descriptions of taxa, follows the Global Cestode 
Database (Caira ef al, 2012). However, the recently 
erected order Onchoproteocephalidea, which groups the 
proteocephalideans and some ‘hooked’ tetraphyllidean 
cestodes (see Caira et al., 2014), is not considered herein 
for the reasons presented by Arredondo et al. (2014), 
especially because no morphological synapomorphies of 
the new order were provided by Caira er al. (2014). Field 
numbers correspond to the numbers of fish examined 
in field protocols (PI = Peru, Iquitos, Loreto Region, 
Peru; letters after host number distinguish different 
worm samples). Since 2008, every fish dissected was 


photographed together with its field number (PI); in 
2009 and 2011, tissue samples, usually a small piece 
of musculature, of every infected fish were taken and 
fixed with 96% molecular-grade ethanol for future DNA 
sequencing to confirm host identification. Photographs of 
fishes and their tissue samples are available upon request 
from the authors. 

Names of teleosts follow those in FishBase (Froese 
& Pauly, 2014) and PlanetCatfish (http://www. 
planetcatfish.com) except for Brachyplatystoma rous- 
seauxii (Castelnau), which was erroneously reported as 
B. flavicans (Castelnau) by de Chambrier et al. (2006a) 
following Fishbase (see Lundberg and Akama, 2005; 
http://www.planetcatfish.com/). In addition, Zungaro 
zungaro (Humboldt) was wrongly reported as Paulicea 
luetkeni (Steindachner) (for current nomenclature, see 
Froese & Pauly, 2014; John Lundberg, pers. comm.). 


RESULTS 


Survey of species found 

Species not reported by de Chambrier ef al. (2006a) 
are marked by an asterisk (*); species reported by de 
Chambrier er al. (2006a), but not found in 2006-2011, are 
also listed herein to provide a complete list of cestodes 
found. Collection numbers refer to the Natural History 
Museum, Geneva, Switzerland — MHNG-PLAT, unless 
otherwise stated. Cestode taxa are listed alphabetically. 


Ageneiella sp.* 


Host: Ageneiosus inermis (L.) (1 fish infected of 16 fish 
examined, i.e. prevalence of 6%). 


Remarks: Specimens found in À. inermis are partly 
decomposed and contracted, which impedes their 
reliable identification to the species level. However, 
they apparently belong to an undescribed species of 
Ageneiella de Chambrier & Vaucher, 1999, a hitherto 
monotypic genus proposed to accommodate A. brevifilis 
de Chambrier & Vaucher, 1999 from Ageneiosus bre- 
vifilis (L.) (synonym of A. inermis) from Paraguay 
by de Chambrier & Vaucher (1999). They possess 
biloculate suckers with a sphincter and lateral lobes of 
the ovary penetrating into the cortex (see de Chambrier 
& Vaucher, 1999). 

Immature cestodes were found in two additional 
Ageneiosus sp., but their identification is not possible. 


Amphoteromorphus ovalis Carfora, de Chambrier & 
Vaucher, 2003* 


Host: Brachyplatystoma cf. filamentosum (Lichten- 
stein); 1/3, 33%). 


Proteocephalidean Cestodes From Peru 151 


Figs 1-5. Scanning electron micrographs of representative scoleces of proteocephalidean cestodes found in the Peruvian Amazon. (1) 
Nomimoscolex lopesi from Pseudoplatystoma fasciatum (PI 708). (2) Proteocephalus sp. 2 from Pterodoras granulosus (PI 
635). (3) Jauella glandicephalus from Zungaro zungaro. (4) Proteocephalus kuyukuyu from Megalodoras uranoscopus (PI 
324). (5) Spatulifer rugosa from P fasciatum (PI 708). 1, 3, 5 = lateral view; 2, 4 = dorsoventral view. 


Amphoteromorphus parkamoo Woodland, 1935 
Host: Zungaro zungaro (3/30, 10%). 


Remarks: Redescribed by Carfora et al. (2003), who 
confirmed the validity of the species. 


Amphoteromorphus peniculus Diesing, 1850* 


Host: Brachyplatystoma rousseauxii (1/3, 33%). 


Amphoteromorphus piriformis Carfora, 
de Chambrier & Vaucher, 2003 


Host: Brachyplatystoma rousseauxii (1/3, 33%). 


Remarks: A. piriformis was described from B. 
rousseauxii collected in Itacoatiara, Brazil in September 
1992 and October 1995 (Carfora et al., 2003). 


Brayela karuatayi (Woodland, 1934)* 
Host: Platynematichthys notatus (Jardine) (5/13, 38%). 


Remarks: This species, which had never been found 
since its original description by Woodland (1934a), has 
recently been redescribed by de Chambrier ef al. (2014) 
on the basis of new material collected in Peru. The 
authors also described correctly its scolex morphology 
and provided the evidence that the actual fish host 
of this cestode is P notatus, not ‘G/anidium sp. as 
reported in the original description (see de Chambrier er 
al., 2014). 


Chambriella agostinhoi (Pavanelli & Machado dos 
Santos, 1992) 


Host: Zungaro zungaro (11/30, i.e. 37%). 


152 A. de Chambrier, R. Kuchta & T. Scholz 


Chambriella paranaensis (Pavanelli & Rego, 1989) 
Host: Hemisorubim platyrhynchos (1/12, 8%). 


Remarks: de Chambrier ef al. (2006a) designated this 
species erroneously as the type species of Chambriella 
Rego, Chubb & Pavanelli, 1999. Rego et al. (1999) 
actually did not explicitly mention the type species 
of the genus in its generic diagnosis on p. 314, but 
C. agostinhoi was mentioned as the type species of the 
genus in remarks to that species (Rego ef al., 1999: 
317), 


Chambriella sp. 1* 


Host: Brachyplatystoma vaillantii (Valenciennes) (4/39, 
10%). 


Remarks: This cestode is a rare parasite of B. vaillantii. 


Chambriella sp. 2 (= Chambriella sp. of 
de Chambrier er al., 2006a) 


Host: Phractocephalus hemioliopterus (Bloch & 
Schneider) (5/10, 50%). 


Chambriella sp. 3* 


Host: Pseudoplatystoma fasciatum (7/42, 17%). 


Chambriella sp. 4* 


Host: Sorubimichthys planiceps (Spix & Agassiz) (6/22, 
27%). 


Remarks: de Chambrier & Scholz (2008) reported but 
did not describe the morphology of this species, which 
may be new to science, similarly as the three species 
listed above (Chambriella spp. 1-3). Their taxonomic 
study is in preparation and will be presented in a 
separate account. 


Choanoscolex abscisus (Riggenbach, 1896) 
Host: Pseudoplatystoma fasciatum (14/42, 33%). 


Remarks: Compared to de Chambrier ef al. (2006a), 
new collections enabled us to obtain sufficient material 
of this cestode, which has been found in a wide 
spectrum of unrelated fish hosts (Rego, 1987, 1990; 
Rego & Pavanelli 1990; Rego er al., 1999). 

In its strobilar morphology, the species closely resembles 
Spatulifer surubim Woodland, 1934 from the same 
fish host, differing only in a much less developed 
metascolex. However, some intermediate forms with a 
more developed metascolex were found, which indicates 
that differences between these taxa of two different 
genera should be critically assessed. Molecular data also 


indicate close relatedness of S. surubim and C. abscisus 
from P. fasciatum (A. Waeschenbach, unpubl. data). 


Choanoscolex sp.* 
Host: Sorubimichthys planiceps (3/22, 14%). 


Remarks: de Chambrier & Scholz (2008) reported 
cestodes of the genus Choanoscolex La Rue, 1911 that 
differ from those of C. abscisus, the only species of the 
genus (see above). 


Endorchis piraeeba Woodland, 1934* 
Host: Brachyplatystoma cf. filamentosum (1/3, 33%). 


Remarks: At present, the genus includes EZ. piraeeba 
from B. filamentosum and Endorchis auchenipteri 
de Chambrier & Vaucher, 1999 from Auchenipterus 
osteomystax (Miranda Ribeiro) from the Parana River in 
Paraguay (de Chambrier & Vaucher, 1999). In addition, 
de Chambrier & Vaucher (1999) reported unidentified 
cestodes of Endorchis from Pimelodus cf. maculatus 
Lacépède and Trachelyopterus striatulus (Steindachner) 
from Paraguay. 

An immature specimen with a similar scolex was found 
in Pseudoplatystoma fasciatum (1/42, i.e. 2%). 


Endorchis sp.* 


Host: Pimelodus altissimus Eigenmann & Pearson 
(1/1). 


Remarks: These specimens resemble those of 
E. auchenipteri but their large-sized Mehlis gland is 
unique among the Proteocephalidea. 


Euzetiella tetraphylliformis de Chambrier, Rego & 
Vaucher, 1999 


Host: Zungaro zungaro (5/30, 17%). 


Remarks: Worms collected in 2009 were immature, 
but are supposed to belong to the only known species 
of the genus, which was described from the same 
host in Itacoatiara, Brazil (de Chambrier et al, 
1999). One immature specimen was also found in 
Pseudoplatystoma fasciatum. 


Gibsoniela mandube (Woodland, 1935)* 


Host: Ageneiosus inermis (2/16, 13%), Ageneiosus sp. 
(3/10, 30%). 


Remarks: The species was described as Anthobothrium 
mandube Woodland, 1935, (Phyllobothriidae) and 
transferred to Gibsoniela Rego, 1984 by Rego (1984). 
From the same host (A. inermis) and same locality 


Proteocephalidean Cestodes From Peru 153 


(Amazon River in Brazil), Woodland (1935a) described 
Endorchis mandube, but Rego (1984) suggested that 
both species may be synonymous and de Chambrier 
(1990) confirmed this synonymy. 

However, de Chambrier & Vaucher (1999) studied the 
type material and newly collected specimens of both taxa 
from the Amazon River and concluded that they represent 
two distinct species of the same genus. To avoid their 
homonymy, they proposed Gibsoniela meursaulti de 
Chambrier & Vaucher, 1999 to accommodate Endorchis 
mandube; tapeworms redescribed by Rego (1992) as 
G. mandube actually belonged to G. meursaulti (de 
Chambrier & Vaucher, 1999). 


Harriscolex kaparari (Woodland, 1935)* 
Host: Pseudoplatystoma fasciatum (3/42, 7%). 


Remarks: Described as Nomimoscolex kaparari 
Woodland, 1935 by Woodland (1935a) from Pseu- 
doplatystoma tigrinum (Valenciennes) in Brazil. 


Harriscolex piramutab (Woodland, 1934) n. comb. 
Host: Brachyplatystoma vaillantii (8/39, 20%). 


Remarks: de Chambrier er al. (2006a) reported 
Proteocephalus piramutab Woodland, 1934 from 
Brachyplatystoma vaillantii. A detailed morphological 
study of newly collected specimens and material of 
Proteocephalus piramutab from museum collections, 
and their comparison with those of AH. kaparari, 
revealed that the former species should be transferred to 
Harriscolex Rego, 1987, because it possesses a scolex 
with a dome-shaped anterior end and suckers with two 
triangular projections (see Rego, 1994). Therefore, a 
new combination, Harriscolex piramutab, is proposed 
for specimens previously identified as P piramutab, 
including those reported from Peru by de Chambrier et 
al. (2006a). 


Houssayela sudobim (Woodland, 1935) 
Host: Pseudoplatystoma fasciatum (3/42, 7%). 


Remarks: Described as Myzophorus  sudobim 
Woodland, 1935 from P fasciatum from the Amazon 
River in Brazil by Woodland (1935b) and found and 
redescribed by de Chambrier & Scholz (2005) for the 
first time since original description on the basis of a 
single specimen from P. fasciatum in Iquitos (PI 76a — 
22. 4. 2004). 


Jauella glandicephalus Rego & Pavanelli, 1985* 
Fig. 3 


Host: Zungaro zungaro (9/30, 30%). 


Remarks: Described from Zungaro jahu (Ihering) (as 
Paulicea luetkeni) from the Parana River in Brazil by 
Rego & Pavanelli (1985). This is the first record of the 
parasite in the Amazon River basin. 


Lenhataenia megacephala (Woodland, 1934)* 
Host: Sorubimichthys planiceps (11/22, 50%). 


Remarks: The genus was erected by de Chambrier 
& Scholz (2008) to accommodate Monticellia mega- 
cephala Woodland, 1934, which is a common, host- 
specific parasite of S. planiceps. 


Manaosia bracodemoca Woodland, 1935* 
Host: Sorubim lima (Bloch & Schneider) (3/29, 10%). 


Remarks: The species was described by Woodland 
(1935a) from ‘Platysoma sp.’ (vernacular name ‘braço 
de moça’) in the Amazon River in Brazil. De Chambrier 
(2003) clarified the systematic position of this species, 
which is a rare parasite of S. lima. He considered 
Paramonticellia Pavanelli & Rego, 1991 to be a junior 
synonym of Manaosia Woodland, 1935. 


Mariauxiella piscatorum de Chambrier & 
Vaucher, 1999 


Host: Hemisorubim platyrhynchos (2/12, 1.e. 17%). 


Remarks: This species was found only in 2004 (de 
Chambrier et al., 2006a). 


Megathylacus jandia Woodland, 1934* 
Host: Zungaro zungaro (2/30, 7%). 


Remarks: This species was originally identified 
as Megathylacus brooksi Rego & Pavanelli, 1985, 
but a detailed study of type and new material of 
Megathylacus cestodes from the Amazon and 
Parana River basins (de Chambrier er al, 2014) 
has demonstrated conspecifity of this species with 
M. jandia, which was described by Woodland (1934a) 
from the Amazon River in Brazil. 


Megathylacus sp.* 
Host: Pseudoplatystoma fasciatum (4/42, 10%). 


Remarks: These cestodes from P fasciatum differ 
from Megathylacus travassosi by a few morphological 
characters and potentially belong to a new species. This 
species was collected also in 2004, but not reported by 
de Chambrier er al. (2006a). 


154 A. de Chambrier, R. Kuchta & T. Scholz 


Monticellia amazonica de Chambrier & Vaucher, 
1997 


Host: Calophysus macropterus (Lichtenstein) (5/33, 
15%). 


Remarks: Scholz er al. (2008) redescribed the species 
on the basis of specimens found in Iquitos, Peru in 2005. 


Monticellia belavistensis Pavanelli, Machado dos 
Santos, Takemoto & dos Santos, 1994* 


Host: Prerodoras granulosus (Valenciennes) (1/24, 4%). 


Remarks: This cestode, which was described by 
Pavanelli et al. (1994) from P granulosus from the 
Parana River basin in Brazil and then reported by de 
Chambrier & Vaucher (1999) from the Paraguay River 
in Paraguay, was found in Peru only once. It is the first 
record of this species from the Amazon River basin. 


Monticellia lenha Woodland, 1933* 
Host: Sorubimichthys planiceps (13/22, 59%). 


Remarks: Originally described by Woodland (1933) 
from specimens found in S. planiceps, and redescribed 
by de Chambrier & Scholz (2008), who studied type 
specimens and new material from Iquitos collected in 
2006 (see Table 2). It is a specific and the most frequent 
parasite of S. planiceps. 


Monticellia santafesina Arredondo & 
Gil de Pertierra, 2010* 


Host: Megalonema platycephalum Eigenmann (1/1). 


Remarks: Described from Megalonema platanum 
(Giinther) from the Parana River basin in Argentina 
(Arredondo & Gil de Pertierra, 2010). This is the first 
geographical record of this cestode from the Amazon 
River basin. M. platycephalum represents a new 
definitive host of the parasite. 


Monticellia ventrei de Chambrier & Vaucher, 1999* 


Host: Pinirampus pirinampu (Spix & Agassiz) (4/30, 
13%). 


Remarks: Monticellia ventrei was described by de 
Chambrier & Vaucher (1999) from specimens found 
in P. pirinampu from the Paraguay River in Paraguay. 
Specimens found in Peru represent a new geographical 
record and expand the distribution area of the species to 
include the Amazon River basin. 


Nomimoscolex admonticellia (Woodland, 1934)* 
Host: Pinirampus pirinampu (11/30, 37%). 


Remarks: This is a relatively common parasite specific 
of P pirinampu, which was originally described by 
Woodland (1934b) from Pinirampus sp. from the 
Amazon River near Itacoatiara, Brazil. 


Nomimoscolex lenha (Woodland, 1933)* 
Host: Sorubimichthys planiceps (5/22, 23%). 


Remarks: de Chambrier & Scholz (2008) redescribed 
the species based on 2 specimens they collected in 
Itacoatiara, Brazil in 1995 and 1 specimen from Iquitos, 
Peru in 2006. Interestingly, the prevalence of N. /enha in 
Brazil (22%, n = 9; see de Chambrier & Scholz, 2008) 
was almost identical to that in the same host from Peru. 


Nomimoscolex lopesi Rego, 1989 
Fig. 1 


Host: Pseudoplatystoma fasciatum (11/42, 26%). 


Remarks: This species was studied using scanning 
electron microscopy for the first time (Fig. 1). 


Nomimoscolex sudobim Woodland, 1935 
Host: Pseudoplatystoma fasciatum (10/42, 24%). 


Remarks: Also found in P tigrinum from Peru by de 
Chambrier et al. (2006a). The species was redescribed 
by de Chambrier er al. (2006b). 


Nomimoscolex suspectus Zehnder, de Chambrier, 
Vaucher & Mariaux, 2000* 


Host: Brachyplatystoma cf. filamentosum (1/3, 33%). 


Remarks: Described from tapeworms found in 
Brachyplatystoma filamentosum (type host), B. flavicans 
(now B. rousseauxii) and B. vaillantii from the Amazon 
River in Brazil (Zehnder et al., 2000); it was found in 
Peru only once. 


Nomimoscolex sp.* 
Host: Pimelodus ornatus Kner (2/13, 15%). 


Remarks: These cestodes differ from N. micro- 
acetabula Gil de Pertierra, 1995 by a few morphological 
characters and potentially belong to a new species. 


Proteocephalidean Cestodes From Peru 155 


Nupelia sp. 


Host: Goeldiella eques (Müller & Troschel) (3/28, 
11%). 


Remarks: Despite great efforts to collect more material 
allowing for its description, only one additional 
specimen from this species was found since 2005 (see 
de Chambrier et al., 2006a). These tapeworms are 
mainly characterized by extraordinarily wide ventral 
osmoregulatory canals. 


Peltidocotyle lenha (Woodland, 1933) 


Hosts: Sorubimichthys planiceps (13/22, 59%); 
Zungaro zungaro (13/30, 43%). 


Remarks: de Chambrier & Scholz (2008) reported 
the species from the type host, S. planiceps. Zungaro 
zungaro is another host of the tapeworm (Zehnder & de 
Chambrier, 2000). 


Peltidocotyle rugosa Diesing, 1850* 
Host: Pseudoplatystoma fasciatum (10/42, 24%). 


Remarks: de Chambrier ef al. (2006a) did not report 
this frequent parasite of P fasciatum. 


Proteocephalus gibsoni Rego & Pavanelli, 1991 
Host: Astronotus ocellatus (1/4, 25%). 


Proteocephalus hemioliopteri de Chambrier & 
Vaucher, 1997* 


Host: Phractocephalus hemioliopterus (1/10, 10%). 


Remarks: de Chambrier & Vaucher (1997) proposed 
a new name, Proteocephalus hemioliopteri, for 
Myzophorus woodlandi Rego, 1984 [syn. Nomimoscolex 
woodlandi (Rego, 1984) Rego & Pavanelli, 1992]; de 
Chambrier er al. (2005) redescribed this species, which 
was found only once in Peru. 


Proteocephalus hobergi de Chambrier & Vaucher, 
1999* 


Host: Oxydoras niger (Valenciennes) (1/16, 6%). 


Remarks: de Chambrier & Vaucher (1999) described 
the species from Oxydoras kneri Bleeker from the 
Parana and Paraguay rivers in Paraguay. The specimens 
found in Peru represent new host and geographical 
records and the first report of the species from the 
Amazon River basin. 


Proteocephalus kuyukuyu Woodland, 1935* 
Fig. 4 


Hosts: Megalodoras uranoscopus (Eigenmann & 
Eigenmann) (3/4, 75%); Pterodoras granulosus (2/24, 
8%); Pterodoras sp. (1/1). 


Remarks: This parasite of doradid catfishes was 
described by Woodland (1935c) from the kuyukuyu, 
vernacular name of Oxydoras niger (as Pseudodoras 
niger), from Codajaz, Brazil. Even though Woodland 
(1935c) found over 50 specimens, no one was mature. 
The same situation was observed in Peru and none of 
347 cestodes found was fully mature. This may indicate 
hyperapolytic development, 1.e. release of proglottids 
precociously before they contain any eggs, which then 
complete their development while free in the intestine of 
the host (see glossary in Khalil er al., 1994), but no free 
proglottids were found in hosts infected with immature 
cestodes. 


Proteocephalus macrophallus (Diesing, 1850) 
Host: Cichla monoculus (2/15, 13%). 


Proteocephalus microscopicus Woodland, 1935 
Host: Cichla monoculus (6/15, 40%). 


Remarks: Both species, similarly as P gibsoni from 
another cichlid (see above), were not found since 2005 
because no other hosts, C. monoculus and A. ocellatus, 
were examined. 


Proteocephalus sophiae de Chambrier 
& Rego, 1994* 


Host: Zungaro zungaro (6/30, 20%). 


Remarks: This is a host-specific parasite of the 
Z. zungaro, described from the Amazon River in Brazil 
by de Chambrier & Rego (1994). 


Proteocephalus sp. 1 of de Chambrier ef al. (2006a) 
Host: Phractocephalus hemioliopterus (2/10, 20%). 


Remarks: Only immature specimens have been found. 


Proteocephalus sp. 2 of de Chambrier ef al., (2006a) 
Fig. 2 


Host: Pterodoras granulosus (2/24, 8%). 


Remarks: Additional immature specimens were found 
in 5 P granulosus and possibly belong to the same 
taxon. 


156 A. de Chambrier, R. Kuchta & T. Scholz 


Proteocephalus sp. 3* 
Host: Pimelodus blochii Valenciennes (2/8, 25%). 


Remarks: All specimens are immature. 


Rudolphiella piracatinga (Woodland, 1935) 
Host: Calophysus macropterus (10/33, 30%). 


Rudolphiella sp.* 
Host: Pinirampus pirinampu (5/30, 17%). 


Remarks: These cestodes differ from both R. myoides 
Woodland, 1934 and R. piranabu Woodland, 1934 from 
the same host in Brazilian Amazon and potentially 
belong to a new species. 


Scholzia emarginata (Diesing, 1850) 
Host: Phractocephalus hemioliopterus (10/10, 100%). 


Remarks: This is the most frequent specific parasite of 
P. hemioliopterus. 


Sciadocephalus megalodiscus Diesing, 1850 
Host: Cichla monoculus (1/15, 7%). 


Remarks: Woodland (1933b) redescribed this taxa 
established by Diesing (1850), based on his material 
collected in the Amazon River in 1931. Rego et al. 
(1999) reported this species from the Parana River basin 
and amended a generic diagnosis. 


Spasskyellina spinulifera (Woodland, 1935) 


Hosts: Pseudoplatystoma fasciatum (12/42, 29%), P. 
tigrinum (Valenciennes) (2/13, 14%; no new material 
found since 2005). 


Remarks: This species was originally described as 
Monticellia spinulifera by Woodland (1935b) from 
P. fasciatum from the Amazon River in Brazil. Freze 
(1965) proposed a new genus, Spasskyellina, to 
accommodate this species. De Chambrier & Vaucher 
(1999) synonymized the genus with Monticellia, but 
later de Chambrier et al. (2006a) listed Spasskyellina 
as a valid genus, ignoring the previous paper from 
1999. Three species of Pseudoplatystoma, namely 
P. corruscans, P. fasciatum and P. tigrinum, from the 
Amazon and Parana River basins in Peru, Brazil and 
Paraguay serve as definitive hosts of S. spinulifera 
(Woodland, 1935b; Rego, 1990; de Chambrier & 
Vaucher, 1999; de Chambrier et al., 2006a; present 
study). 


Spatulifer maringaensis Pavanelli & Rego, 1989 


Host: Hemisorubim platyrhynchos (Valenciennes) 
(2/12; 17%); Sorubim lima (3/29, 10%). 


Remarks: Originally described from A. platyrhynchos 
and found by de Chambrier er al. (2006a) in the same 
host from Peru. Arredondo & Gil de Pertierra (2008) 
confirmed that tapeworms from this catfish and S. lima 
from the Parana River basin are conspecific, which was 
supported by the present study. 


Spatulifer rugosa (Woodland, 1935) 
Fig. 5 


Host: Pseudoplatystoma fasciatum (14/42, 33%). 


Remarks: Described as Monticellia rugosa from 
P. fasciatum from the Amazon River, Brazil by 
Woodland (1935a), who reported prevalence of 55%. 


Spatulifer sp. (probably S. surubim Woodland, 1934) 
Host: Pseudoplatystoma tigrinum (1/13, 8%). 


Remarks: Only immature specimens were found; see 
de Chambrier ef al. (2006a). 


Travassiella jandia (Woodland, 1934) 
Host: Zungaro zungaro (1/30, 3%). 


Remarks: de Chambrier et al. (2014) synonymized 
Travassiella avitellina Rego & Pavanelli, 1987 
described from Zungaro zungaro (in fact Zungaro jahu, 
see Lundberg & Littmann, 2003) with Proteocephalus 


jandia Woodland, 1934 described from Z. zungaro 


from the Amazon River in Brazil and proposed a new 
combination, 7. jandia (Woodland, 1934). This is the 
rarest species found in Z. zungaro in Peru. 

No new specimens were found since 2004 (see de 
Chambrier et al., 2006a). 


Zygobothrium megacephalum Diesing, 1850 
Host: Phractocephalus hemioliopterus (1/10, 10%). 


Monticelliinae gen. sp.* 
Host: Phractocephalus hemioliopterus (1/10, 10%). 


Remarks: This material differs from all known species 
of the Monticelliinae and potentially belong to a new 
species and genus. 


Proteocephalidean Cestodes From Peru 157 


Proteocephalidea gen. sp.* 
Host: Cichlasoma amazonarum Kullander (3/29, 10%). 


Remarks: This material differs from all known species 
of the Proteocephalidae and potentially belong to a new 
species and genus. 


DISCUSSION 


Our four recent sampling trips in the Peruvian Amazon 
enable us to double the number of proteocephalidean 
cestodes reported in 25 species of fishes of the upper 
part of the Amazon River around Iquitos. Besides the 
29 proteocephalidean species found in 8 species of 
pimelodid, 1 heptapterid and 1 doradid catfishes, and 2 
species of cichlids, the present account adds another 34 
species, thus providing evidence of the occurrence of as 
many as 63 species of these cestodes in Peru. Out of them, 
46 species could be identified to the species level and 
represent already known taxa. This is only slightly less 
than the known fauna of the Brazilian part of Amazonia, 
from which 54 species have been reported (Table 1). 
However, more than a dozen species found only in Peru 
are probably new taxa awaiting formal description. In 
total, as many as 64 named species of proteocephalidean 
cestodes are now known from the Amazon River basin 
(Table 1). 

Unlike Peru, with almost no history of studies on cestode 
parasites of freshwater teleosts, research on fish parasites 
in the Brazilian part of the Amazon River basin started 
as early as in the 19th century, when Diesing (1850) 
described several taxa based on material collected by 
an Austrian naturalist and explorer Johann Natterer, 
who spent 18 years in Brazil from 1817 to 1835. More 
systematic studies on proteocephalidean cestodes started 
in the early 1930’s when W.N.F. Woodland published 
9 papers with descriptions of 32 species and 8 genera 
of proteocephalidean cestodes from catfishes and 
other teleost fishes in the Amazon River in Brazil (de 
Chambrier et al., 2014). After a couple of decades since 
Woodland’s pioneer studies, Brazilian authors, especially 
A.A. Rego, studied the cestode fauna of fishes in Brazil 
(see Rego et al., 1999 for references), even though many 
of the studies were carried out in the Parana River basin 
(Rego & Pavanelli, 1992; Fig. 6). Since the mid 1990’s, 
the senior author with co-authors have also contributed 
considerably, with a number of papers dealing with 
proteocephalideans from the Brazilian Amazon (see de 
Chambrier et al., 2006a, 2014 and references therein). 
From the Amazon River basin in Peru, de Chambrier 
et al. (2006a) reported 5 species of proteocephalideans 
from Pseudoplatystoma fasciatum and Paulicea 
luetkeni (= Zungaro zungaro) each, and 4 species in 
Phractocephalus hemioliopterus and Pseudoplatystoma 
tigrinum. In the present study, much higher numbers of 
cestodes were found in these hosts: 10 (and juveniles of 


2 other species) in P fasciatum (plus one unidentified 
species; 7 of these species were also reported from 
Brazilian Amazonia), 9 in Z. zungaro (1 unidentified 
species) and 6 in P hemioliopterus (1 unidentified 
species). As typical for proteocephalidean cestodes in 
the Neotropical Region (e.g., de Chambrier & Vaucher, 
1999), most species are specific to a single fish host, 
more rarely to congeneric host species. In the present 
study, 9 species were found in more than one fish host 
and only 2 species, namely Proteocephalus kuyukuyu 
and Spatulifer maringaensis, occur in fish of more than 
one genus (Table 2). However, there could be a sampling 
bias, considering that the number of dissected hosts per 
species varied considerably, from 1 to 42 specimens 
(Table 2). Pseudoplatystoma fasciatum Was the most 
frequently examined host and also harboured the highest 
number of species of proteocephalideans. 

Extensive material of proteocephalidean cestodes was 
collected in a wide spectrum of teleosts during six 
visits by the present authors and their co-workers to the 
Peruvian Amazonia. This material will make it possible 
to compare the species composition of the cestode fauna 
and host-parasite associations in the Amazon River 
basin with those in the Parana River basin (Fig. 6). 
Some proteocephalideans occur in closely related hosts 
from different river basins, such as Proteocephalus 
macrophallus and P. microscopicus in species of Cichla 
Bloch & Schneider, 1801, or in recently separated 
‘couples’ of fish hosts that occur only in one of the two 
principal river basins in South America, 1.e. Amazon 
and Parana, such as Zungaro zungaro in the former river 
basin and Z. jahu in the latter one. 


IN BO ® Rio Pde Janeiro 
I =” Sao Paulo 


fn re 
“Se we \ 
de Sx 
Fe ie > 
{ 


Fig. 6. Map of South America with two principal river basins, 
Amazon and Parana Rivers. 


158 A. de Chambrier, R. Kuchta & T. Scholz 


Table 1. List of species of proteocephalidean cestodes found in fishes from the Amazon River basin. 


Species 

Amazotaenia yvettae de Chambrier, 2001 
Amphoteromorphus ninoi Carfora, de Chambrier & Vaucher, 2003 
Amphoteromorphus ovalis Carfora, de Chambrier & Vaucher, 2003 
Amphoteromorphus parkamoo Woodland, 1935 
Amphoteromorphus peniculus Diesing, 1850 
Amphoteromorphus piraeeba Woodland, 1934 
Amphoteromorphus piriformis Carfora, de Chambrier & Vaucher, 2003 
Brayela karuatayi (Woodland, 1934) 

Brooksiella praeputialis (Rego, dos Santos & Silva, 1974) 
Chambriella agostinhoi (Pavanelli & Machado dos Santos, 1992) 
Chambriella paranaensis (Pavanelli & Rego, 1989) 
Choanoscolex abscisus (Riggenbach, 1896) 

Endorchis piraeeba Woodland, 1934 

Ephedrocephalus microcephalus Diesing, 1850 

Euzetiella tetraphylliformis de Chambrier, Rego & Vaucher, 1999 
Gibsoniela mandube (Woodland, 1935) 

Gibsoniela meursaulti de Chambrier & Vaucher, 1999 
Goezeella siluri Fuhrmann, 1915 

Harriscolex kaparari (Woodland, 1935) 

Harriscolex piramutab (Woodland, 1934) n. comb. 
Houssayela sudobim (Woodland, 1935) 

Jauella glandicephalus Rego & Pavanelli, 1985 

Lenhataenia megacephala (Woodland, 1934) 

Mariauxiella piscatorum de Chambrier & Vaucher, 1999 
Manaosia bracodemoca Woodland, 1935 

Megathylacus jandia Woodland, 1934 

Monticellia amazonica de Chambrier & Vaucher, 1997 
Monticellia belavistensis Pavanelli et al., 1994* 

Monticellia lenha Woodland, 1933 

Monticellia magna (Rego, Santos & Silva, 1974) 

Monticellia santafesina Arredondo & Gil de Pertierra, 2010 
Monticellia ventrei de Chambrier & Vaucher, 2009 
Nomimoscolex admonticellia (Woodland, 1934) 
Nomimoscolex dorad (Woodland, 1935) 

Nomimoscolex lenha (Woodland, 1933) 

Nomimoscolex lopesi Rego, 1989 

Nomimoscolex microacetabula Gil de Pertierra, 1995 
Nomimoscolex piraeeba Woodland, 1934 

Nomimoscolex sudobim Woodland, 1935 

Nomimoscolex suspectus Zehnder et al., 2000** 

Nupelia portoriquensis Pavanelli & Rego, 1991 
Peltidocotyle lenha (Woodland, 1933) 

Peltidocotyle rugosa Diesing, 1850 

Proteocephalus gibsoni Rego & Pavanelli, 1991 
Proteocephalus hemioliopteri de Chambrier & Vaucher, 1997 
Proteocephalus hobergi de Chambrier & Vaucher, 1999 
Proteocephalus kuvukuyu Woodland, 1935 

Proteocephalus macrophallus Diesing, 1850 

Proteocephalus microscopicus Woodland, 1935 
Proteocephalus platystomi Lynsdale, 1959 

Proteocephalus sophiae de Chambrier & Rego, 1994 
Pseudocrepidobothrium eirasi (Rego & de Chambrier, 1995) 
Pseudocrepidobothrium ludovici Ruedi & de Chambrier, 2012 
Rudolphiella myoides (Woodland, 1934) 

Rudolphiella piracatinga (Woodland, 1935) 

Rudolphiella piranabu (Woodland, 1934) 

Scholzia emarginata (Diesing, 1850) 

Sciadocephalus megalodiscus Diesing, 1850 

Spasskyellina spinulifera (Woodland, 1935) 

Spatulifer maringaensis Pavanelli & Rego, 1989 

Spatulifer rugosa (Woodland, 1935) 

Spatulifer surubim Woodland, 1934 

Travassiella jandia (Woodland, 1934) 

Zygobothrium megacephalum Diesing, 1850 


Total 64 


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* Pavanelli, Machado dos Santos, Takemoto & dos Santos, 1994 
**Zehnder, de Chambrier, Vaucher & Mariaux, 2000 


159 


Proteocephalidean Cestodes From Peru 


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Proteocephalidean Cestodes From Peru 161 


In the present study, we focused on adult cestodes 
and thus only very few proteocephalidean larvae 
(merocercoids, see Chervy, 2002) were found. The 
identification of the latter based on morphological 
characteristics is impossible, but their molecular analysis 
will enable us to match their DNA sequences with those 
of adults, which were obtained in the framework of a 
NSF-PBI project “A Survey of the Tapeworms (Cestoda: 
Platyhelminthes) from the Vertebrate Bowels of the 
Earth” (www.tapeworms.uconn.edu). This approach (see 
also Jensen & Bullard, 2010) is the most feasible way to 
elucidate life cycles of Neotropical proteocephalideans, 
which are almost completely unknown. Based on the fact 
that large catfishes are predatory (piscivorous) and do 
not consume plankton, it is reasonable to assume that life 
cycles of many, if not most, proteocephalidean cestodes 
that mature in large catfishes in South America include 
two intermediate (or one intermediate and one paratenic) 
hosts, the second host being small planktonophagous fish 
similarly to the developmental cycles of Proteocephalus 
ambloplitis in North America (Hunter, 1928; Freze, 1965; 
Scholz, 1999; Scholz & de Chambrier, 2003). 

To summarize, the present study enriched considerably 
the current knowledge of the species composition and 
distribution areas of members of one of the dominant 
groups of metazoan parasites of freshwater teleosts in the 
Neotropical Region. However, there are still many fish 
hosts that have never been examined for parasites and thus 
a number of new taxa probably remain to be discovered. 
This will depend on sampling effort and time dedicated to 
future collecting trips and taxonomic evaluation of newly 
collected cestodes, using combined morphological and 
molecular approaches. A recent discovery of a new genus 
from the little known auchenopterid catfish Tocantinsia 
piresi from the Xingu River in Brazil (Alves et al., 2015) 
as well as the number of unidentified species found in 
Peru, which may represent new species, supports the 
argument that we are just at the very beginning of a 
long path towards representative mapping the species 
diversity, host-parasite relationships and zoogeography 
of the parasites of teleost fishes in one of the hottest 
spots of fish parasite diversity on the Earth. Since the 
existence of cryptic species among proteocephalidean 
cestodes cannot be excluded, it is impossible to provide 
a reliable estimate of the species richness of these 
parasites. However, it is reasonable to assume that the 
actual number of extant species would be at least twice 
as high as the number of the species currently considered 
to be valid. 


ACKNOWLEDGEMENTS 


The authors are deeply indebted to Jean Mariaux, Florian 
Reyda and Amilcar Arandas Rego for helpful suggestions, 
to Martin Mortenthaler and his family, Acuario Rio 
Momon, Iquitos, Peru, for providing facilities for fish 


examination and supplying fish from Santa Clara, Rio 
Nanay. Sophie de Chambrier, Sandrine C. Coquille 
and Carlos A. Mendoza-Palmero helped with fish 
examination in 2009 and 2011. Thanks are due to John G. 
Lundberg, Department of Ichthyology, The Academy of 
Natural Sciences, Philadelphia, Pennsylvania, USA, for 
valuable remarks on the identification of some fish hosts, 
and to Andrea Waeschenbach, Natural History Museum, 
London, UK, for providing unpublished data on the 
molecular phylogeny of proteocephalidean cestodes. 
Support of authorities of the Czech Embassy in Lima, 
Peru and Peruvian Embassy in Prague, Czech Republic, 
in particular of Marina Landaveri, Ambassador of Peru 
in the Czech Republic, is also greatly appreciated. The 
stays of the present authors in Peru were financially 
supported by the National Science Foundation, USA 
(PBI award Nos. 0818696 and 0818823), Czech Science 
Foundation (project No. P505/12/G112) and the Institute 
of Parasitology, AS CR, Ceské Budëjovice (RVO: 
60077344). 


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À 


iR 


Revue suisse de Zoologie (March 2015) 122(1): 165-180 


ISSN 0035-418 


Review of the Himalayan genus Hingstoniella Jeannel, 
and description of Besuchetaceus gen. n. from Nepal 
(Coleoptera: Staphylinidae: Pselaphinae) 


Zi-Wei Yin & Li-Zhen Li 


Department of Biology, College of Life and Environmental Sciences, Shanghai Normal University, 100 Guilin Road, 


Shanghai, 200234, P. R. China 


* Corresponding author. E-mail: pselaphinae@gmail.com 


Abstract: The Himalayan genus Hingstoniella Jeannel and its type species A. lata Jeannel are redescribed, and a new 
species, H. trigona sp. n., is described. Besuchetaceus gen. n. is established for a single species, B. nepalensis sp. n., from 
central Nepal. Both genera together with Sinotrisus Yin & Li are placed in the newly designated ‘Hingstoniella group’. 
Keys are provided to distinguish genera of Hingstoniella group and species of Hingstoniella, and the major diagnostic 


features of all included taxa are illustrated. 


Keywords: Taxonomy - Batrisitae - Hingstoniella - Besuchetaceus - Sinotrisus - new genus - new species - Himalayan 


region. 


INTRODUCTION 


In his study of the northern Indian Pselaphinae, Jeannel 
(1960) created Hingstoniella Jeannel for a new species, 
H. lata Jeannel, on the basis of one male from Sikkim 
and one female from Yatung, Tibet (= Yadong, Xizang 
A. R.). Jeannel stated (1960: 410) that Hingstoniella 
was ‘of uncertain affinity, but however appearing to 
be well related to Indo-Malaysian lineages’. Based on 
the additional material from Tibet, Yin ef al. (2011) 
redescribed the genus and species, and reported its 
association with the ant genus Myrmica Latreille. 

The discovery and a subsequent revision (Yin et al. 
2010, 2012) of the genus Sinotrisus Yin & Li, with four 
species scattered in China and Vietnam, seemed to shed 
some light on the uncertain relationship of Hingstoniella 
in Batrisina. Despite the quite different shape of the 
pronotum, both genera share a modified vertex in male, 
and a considerably constricted base of the aedeagus — a 
type of aedeagus that is unique in Batrisitae, which is 
considered to be a possible synapomorphy of these two 
genera. 

As part of our on-going study of the on loan MHNG 
material, in this paper we redescribe Hingstoniella and 
H. lata, document many new locality records for the type 
species in Nepal and neighboring area, and describe a new 
species, H. trigona sp. n., also from Nepal. In addition, 
a new genus and species, Besuchetaceus nepalensis gen. 
et sp. n., is described from Nepal. Since Besuchetaceus 
shares with Hingstoniella and Sinotrisus the modified 


Manuscript accepted 17.11.2014 
DOI: 10.5281/zenodo. 14576 


male vertex and the unique type of aedeagus, these taxa 
are placed into a small group of genera here termed the 
‘Hingstoniella group’. Identification keys to genera of 
Hingstoniella group as well as to species of Hingstoniella 
are provided, and diagnostic features of all included taxa 
are illustrated. 


MATERIAL AND METHODS 


Material used in this paper are housed in the following 
public institutions: 

MHNG Muséum d’ Histoire Naturelle, Geneva, Switzer- 

land (Giulio Cuccodoro) 
SNUC Insect Collection of the Shanghai Normal 
University, Shanghai, China (Zi-Wei Yin) 

The collection data of the material are quoted verbatim. A 
slash (/) is used to separate different labels, authors’ notes 
are included in square brackets ([]). The terminology 
follows Chandler (2001), except we use ‘ventrite’ instead 
of ‘sternite’ when describing meso- and metathoracic 
structures; the terms ‘tergite’ and ‘sternite’ are used when 
referring to abdominal segments. 

The following abbreviations are applied: AL - length of 
the abdomen along the midline; AW - maximum width 
of the abdomen; EL - length of the elytra along the 
sutural line; EW - maximum width of the elytra; HL - 
length of the head from the anterior clypeal margin to 
the occipital constriction; HW - width of the head across 
eyes; PL - length of the pronotum along the midline; PW 


166 Z.-W. Yin & L.-Z. Li 


- maximum width of the pronotum. The total body length 
is a combination of HL + PL + EL + AL. 


TAXONOMY 
Hingstoniella group of genera 


Definition: Members of the Hingstoniella group have a 
more-or-less-developed modified vertex in the male, and 
share a strongly constricted base of the aedeagus, this 
type of aedeagus seems to be unique in Batrisitae. 


Included taxa: Hingstoniella, Sinotrisus, and Besu- 
chetaceus gen. n. 


Hingstoniella Jeannel, 1960 
Figs 1-3 
Hingstoniella Jeannel, 1960: 410. Type species: Hingstoniella 
lata Jeannel (original monotypy). 
Hingstoniella: Newton & Chandler, 1989: 37 (catalog); 
Yin, Li & Zhao, 2011: 390 (diagnosis, redescription, 
illustrations). 


Diagnosis: Head triangular; frontal rostrum low; with 
punctiform vertexal foveae; antennomeres XI elongate 
and conical. Pronotum transversely octagonal, with only 
punctiform lateral antebasal foveae; lacking antebasal 
spines. Elytra each with one large basal fovea and 
shallow discal stria. Abdomen with lateral margins of 
tergite IV edged by marginal carinae; tergite IV longest. 


Redescription: General body form robust; Length 3.09- 
3.56 mm. Head triangular (Fig. 1A); with low frontal 
rostrum, antennal tubercles low; vertexal foveae small 
and punctiform, connected by indistinct U-shaped 


Key to genera of Hingstoniella group 


vertexal sulcus; antennae with 11 antennomeres, clubs 
formed by apical three antennomeres, antennomeres 
XI elongate and conical; ocular-mandibular carinae 
faint; eyes reniform (Fig. 1B); maxillary palpi with 
palpomeres III transverse and triangular, IV fusiform; 
with gular foveae merged into single opening (Fig. 1C). 
Pronotum (Fig. 1D) transversely octagonal; lateral 
antebasal foveae punctiform, lacking median antebasal 
fovea and antebasal spines; disc only slightly convex; 
lacking basolateral foveae; lacking paranotal carinae; 
lateral procoxal foveae (Fig. 1E) present. Each elytron 
(Fig. IF) with one large basal fovea, discal striae 
shallow, sutural striae complete; lacking subhumeral 
foveae, marginal stria present from basal one-fourth 
toward posterior margin; lateroapical notch indistinct. 
Thorax (Fig. 2A) with small median and lateral 
mesoventral foveae; with lateral mesocoxal foveae; 
prepectal foveae present; lateral metaventral foveae 
punctiform, metacoxae narrowly separated; posterior 
margin with narrow median notch. Abdomen (Fig. 
2B-D) has tergites IV-VI (first to third visible tergites) 
with marginal carinae extending through tergal length; 
tergite IV longest, V-VII slightly shorter and subequal 
in length; tergite IV with basolateral foveae at lateral 
ends of sulcus, lacking mediobasal foveae and discal 
carinae, V-VII each with one pair of basolateral foveae. 
Sternite IV (second visible ventrite) twice length of V at 
midlength, with one pair of mediobasal foveae and three 
pairs of basolateral foveae, lacking basal sulcus, V-VII 
lacking foveae. Legs short, second and third tarsomeres 
subequal in length. 

Male (Fig. 3A-B) with vertex modified. Aedeagus with 
paramere fused to median lobe to form elongate ventral 
lobe; articulated dorsal lobe present; basal capsule with 
strongly constricted base. 


| 


Nw 


General form stouter (Fig. 3); pronotum distinctly transverse (Figs 1D, 1E, 3); each elytron with one basal fovea 
(Fig: TP esse TR D SR A RU RE Hingstoniella Jeannel 
General form slenderer (Yin ef al., 2010: fig. 2; Yin et al., 2012: figs 1, 4; Fig. 10); pronotum slightly transverse 
to slightly longer than wide; each elytron with three basal foveae (Yin ef al., 2012: figs 2A, 3A, SA, 6A; Figs 8F, 


Pronotum less transverse, lateral margins lacking spines or denticles (Yin et al., 2012: figs 2A, 3A, SA, 6A); deep 
basal sulcus of tergite IV interrupted by discal carinae or ridges (Yin et al., 2012: figs 1, 4) .. Sinotrisus Yin & Li 
Pronotum more transverse, lateral margins spinose or dentate (Fig. 8D); basal sulcus of tergite IV shallow, lacking 
discallcarinaeto rid esi (E12S19 CR) OI PR Besuchetaceus gen. n. 


Key to males of Hingstoniella 


I 


Vertexal modification composed of indistinct punctiform protuberance (Fig. 4A); each eye with more than 60 
facets; protibiae with mesal margins angularly expanded near apex (Fig. 4C). Distribution: Nepal, Tibet, northern 
India (Fig: 12). ni RE n: H. lata Jeannel 
Vertexal modification composed of distinct triangular bump (Fig. 7A); each eye with about 40 facets; protibiae 
with mesal margins less expanded (Fig. 7C). Distribution: Nepal (Fig. 12).............. H. trigona sp. n. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 167 


Comparative notes: The genus up on which the 
Hingstoniella group is based. Members of the group 
share a modified vertex in the male, and a strongly 
constricted base of the aedeagal basal capsule. 
Hingstoniella can be quickly separated from Sinotrisus 


and Besuchetaceus gen. n. by the transversely octagonal 
pronotum, and each elytron with single basal fovea, 
while both latter genera have the pronotum slightly 
transverse or slightly longer than wide, and have three 
basal foveae on each elytron. 


Fig. 1. Morphology of Hingstoniella. (A) Head, in dorsal view. (B) Same, in lateral view. (C) Same, in ventral view. (D) Pronotum. 
(E) Prosternite. (F) Left elytron. Abbreviations: bef - basal elytral fovea; ds - discal striae; gf - gular foveae; laf - lateral 
antebasal foveae; Ipcf - lateral procoxal foveae; mls - median longitudinal sulcus; ss - sutural striae; vf - vertexal foveae. 


Scales: 0.2 mm. 


168 Z.-W. Yin & L.-Z. Li 


Fig. 2. Morphology of Hingstoniella. (A) Meso- and metaventrite. (B) Abdomen, in lateral view. (C) Same, in dorsal view. (D) Same, 
in ventral view. Abbreviations: blf - basolateral foveae; bs - basal sulcus; Imcf - lateral mesocoxal foveae; Imsf - lateral 
mesoventral foveae; Imtf - lateral metaventral foveae; mbf - mediobasal foveae; mmsf - median mesoventral foveae; ppf - 
prepectal foveae. Scales: 0.5 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 169 


Hingstoniella lata Jeannel, 1960 
Figs 3A, 4-6, 12 
Hingstoniella lata Jeannel, 1960: 410. Type locality: Sikkim, 
Lachen, alt. 3000 m. 


Hingstoniella lata. — Yin et al., 2011: 395 (description, new 
locality, host record). 


Material examined: (23 44, 23 99). CHINA (in 
SNUC): 3 GG, 1 9, labeled ‘China: Xizang A. R., 
Neilamu Co., Quxiang (HZ), alt. 3300 m, nest of 
Myrmica sp. under rock, 20.v11.2010, Wen-Xuan Bi 
leg. [1 ex Myrmica each pinned under two males]’. — 
1 3, 1 ©, labeled ‘China: Xizang A. R., Yadong Co., 
Xiayadong ( FE), alt., 3400 m, nest of Myrmica 
sp. under rock, 9.viii.2010, Wen-Xuan Bi leg. [1 ex 
Myrmica pinned under a male]’. — 2 44, 2 99, same 
locality, except 8.vi11.2010, 2800 m [1 ex Myrmica 
pinned under a male]’. - NEPAL (in MHNG): 1 4, 
labeled ‘Thodung via, Those 3100 m, 29.-31.5.1976 
/ Nepal, W. Wittmer, C. Baroni Urbani / comparé 
au type / Hingstoniella lata Jeann. Cl. Besuchet 
dét. XII. 1977’. — 1 ©, labeled ‘Pokhara 820 m, 15.- 
18.6.1976 / Nepal, W. Wittmer, C. Baroni Urbani / 
Hingstoniella lata Jeann. Cl. Besuchet dét. XII. 1977’. 
— 1 6, labeled ‘NEPAL (Pro. Bagmati), Ridge betw. 
Mere Dara and Thare Pati, 3500 m, Lôbl & Smetana, 
9.IV.81 [2 ex. Myrmica pinned under specimen]’. — 
1 3, labeled ‘Umg, Alm Dugdinma, b. Lughla, 3000- 
4000 m, Khumbu, Nepal, lg. Franz [Pa263, overleaf] / 
Hingstoniella, det. D. S. Chandler’. — 5 © © [on three 
pins, each pin also with 2x Myrmica], labeled ‘NEPAL 
(Prov. Bagmati), below Jangtang Ridge, NE Barahbise, 
3150 m, Lôbl & Smetana, 4.V.81 / Hingstoniella’. 
— 1 ©, labeled ‘NEPAL, Khumbu, Periche 4350 m, 
29.vi.1963, leg. G. Ebert’. — 1 ©, labeled ‘NEPAL: 
Bagmati, Pokhare NE Barahbise, 2800 m, 3.5.81, Löbl 
- Smetana’. — 3 GG, 2 29, labeled ‘NEPAL, 9.x.84, 
Gairi, L. Deharveng’ [according to pers. comm. with 
L. Deharveng, detail collection data of this material is 
as following: ‘Nepal: Trekking of Namche Bazar from 
le Tomba-Kosi: Massif of Hanumante Danda above Jiri. 
Lieu-dit Gairi, 09/10/84, forest, litter, by hand, Louis 
Deharveng leg (NEP84-04)’]. — 1 ©, labeled ‘Gufa- 
Gorza, 2800-2100 m, 4.VI.1985 / E-Nepal, Koshi, 
M. Brancucci’. — 1 ©, labeled ‘E-NEPAL, Dhankuta, 
Arun Valley, SE des Makalu, Tashigaon, 13.-14.6.1980, 
leg. C. Holzschuh, 2100 m / 195 ’. — 3 Sg, labeled ‘E 
Nepal, Koshi, Chauki, 2700 m, Holzschuh, 2.vi.85° [1 
ex Myrmica each pinned under 2 males, 2 ex Myrmica 
under the other male]. - 2 44, 2 29, labeled ‘“Bakan” 
W of Tashigaon 3200 m, 6.1V.1982, A. & Z. Smetana / 
NEPAL, Khandbari District’ [1 ex Myrmica each pinned 
under a male and a female]. — INDIA (in MHNG): 4 
SS, 4 29, labeled ‘INDIA: Darjeeling distr., Tongli, 
3100 m, 16.x.1978, I, Löbl, Bes., nr. 16a, under stones’ 
[1 ex Myrmica each pinned under two males and one 


female]. - 2 Sg, 1 ©, labeled ‘INDIA - West Bengal 
Distr. Darjeeling, Tonglu, 16.X.78, Besuchet- Löbl’. 


Each of the above specimens bears an identification 
label as ‘Hingstoniella lata Jeannel, 1960, det. Yin & Li, 
2014. 


Description: Male (Fig. 3A). BL 3.20-3.41 mm. Body 
reddish brown; most part of dorsal surface densely 
hairy. 

Head (Fig. 4A) distinctly transverse, HL 0.55-0.59 mm, 
HW 0.74-0.84 mm; vertex slightly convex, modification 
composed of indistinct punctiform protuberance, 
foveae located below level of posterior margins of 
eyes; each eye with about 65 small facets; antennal 
clubs formed by apical three enlarged antennomeres 
(Fig. 4B). Pronotum transversely octagonal, PL 0.65- 
0.74 mm, PW 1.02-1.09 mm, median longitudinal 
sulcus weakly developed. Elytra wider than long, EL 
1.02-1.03 mm, EW 1.28-1.31 mm; shallow discal 
striae extending to 3/4 of elytral length. Metathoracic 
wings fully developed. Protibiae (Fig. 4C) triangularly 
projected at mesal margins near apices; mesotrochanters 
(Fig. 4D) with small protuberance at ventral margin; 
metatrochanters (Fig. 4E) with ventral margin 
protruding to form strong projection. Abdomen wider 
than long, AL 0.98-1.05 mm, AW 1.17-1.20 mm. 
Aedeagus (Fig. 4F-I) asymmetric, length 0.49-0.52 mm. 
Female. Similar to male in general appearance, 
protibiae, mesotrochanters, and metatrochanters lacking 
protuberance or projection. Each eye with about 45 
facets. Tergite VIII (Fig. 4J) and sternite VIII (Fig. 4K) 
transverse. Genital complex (Fig. 4L) transverse, width 
0.37 mm. Measurements: BL 3.09-3.29 mm, HL 0.52- 
0.55 mm, HW 0.75-0.81 mm, PL 0.66-0.72 mm, PW 
1.06-1.08 mm, EL 0.93-1.00 mm, EW 1.06-1.20 mm, 
AL 0.98-1.02 mm, AW 1.15-1.17 mm. 


Differential diagnosis: Hingstoniella lata can be 
separated from A. trigona sp. n. described below only 
by the male vertexal modification with an indistinct 
punctiform protuberance; in contrast, the male of 
H. trigona has the vertexal modification represented by 
a much larger, triangular bump. 


Intraspecific variation: While the male vertexal 
modification and other characters on legs are stable, the 
form of aedeagus of populations from different localities 
vary distinctly, especially the dorsal lobe in axial view. 
When available, at least one male from each locality 
was dissected to illustrate the intraspecific variation 
(Figs 4F-I, 5, 6). 


Biology: Adults of this species were frequently 
collected from nests of Myrmica ants nesting under 
rocks. 


Distribution: Nepal: Gandaki, Bagmati, Janakpur, 
Sagarmatha, Koshi; India: Sikkim, West Bengal; China: 
Xizang (Fig. 12). 


170 Z.-W. Yin & L.-Z. Li 


Hingstoniella trigona sp. n. 
Figs SB;7112 


Holotype: Holotype 4, NEPAL: labeled ‘Taksanggeb., 
b. Tukche, Takola [Pa327-327a, overleaf] / Zentral- 
Nepal, Sept.-Okt. 1971, lg. H. Franz / n. sp. / 
HOLOTYPE [red], ©, Hingstoniella trigona sp. n., det. 
Yin & Li, 2014, MHNG’; in MHNG, without accession 
number. 


Paratypes: NEPAL, 1 £, labeled ‘NEPAL, Kali 
Gandaki, vall., Nilgiri Himal. upp., Titigaon vill., 3000- 
3200 m NN, 21.V.2002, leg. J. Schmidt’. — 1 ©, labeled 
‘NEPAL, Annapurna Reg., Umg. Ghorepani, 2800 m, 
28°24'15"N, 83°42'01"E, 22.1V.2000, leg. A. Weigel’. — 
1 ©, labeled ‘Zentral - Nepal, Annapurna - SW, 16.4.82, 
4500 m, Baum / COLL. PANKOW [1 ex Myrmica 
pinned under specimen]’; in MHNG, without accession 
number. 


Each paratype bears a type label similar to that of 
holotype except ‘Paratype [yellow], & [or 9]?. 


Description: Male (Fig. 3B). BL 3.36-3.44 mm. Body 
reddish brown; most part of dorsal surface densely 
hairy. 

Head (Fig. 7A) distinctly transverse, HL 0.58-0.60 mm, 
HW 0.85-0.86 mm; vertex slightly convex, modification 
composed of distinct triangular bump, foveae located 
below level of posterior margins of eyes; each eye with 
about 40 small facets; antennal clubs formed by apical 
three enlarged antennomeres (Fig. 7B). Pronotum 
transversely octagonal, PL 0.70-0.73 mm, PW 1.04- 
1.06 mm, median longitudinal sulcus moderately 
developed. Elytra wider than long, EL 1.02-1.04 mm, 
EW 1.26-1.30 mm; shallow discal striae extending to half 
of elytral length. Metathoracic wings fully developed. 
Protibiae (Fig. 7C) slightly projected at mesal margins 
near apices; mesotrochanters (Fig. 7D) with small 


Fig. 3. Habitus of Hingstoniella. (A) H. lata. (B) H. trigona Scales: 0.5 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 171 


Fig. 4. Diagnostic characters of Hingstoniella lata (A-I Male, J-L Female.). (A) Head and pronotum. (B) Antenna. (C) Apical portion 
of protibia. (D) Mesotrochanter. (E) Metatrochanter. (F) Aedeagus of population from Xiayadong, Tibet, in axial view. (G) 
Same, in dorsal view. (H) Same, in lateral view. (1) Same, in ventral view. (J) Tergite VIII. (K) Sternite VII. (L) Genital 
complex. Scales: A, B = 0.5 mm; F-L = 0.2 mm; C-E = 0.1 mm. 


172 Z.-W. Yin & L.-Z. Li 


Fig. 5. Aedeagus of Hingstoniella lata in populations from various localities (A, E, I, M Dorsal view. B, F, J, N Lateral view. C, G, K, 
O Ventral view. D, H, L, P Axial view.). (A-D) Population from Tonglu, Darjeeling, India. (E-H) Population from “Bakan” W 
of Tashigaon, Nepal. (I-L) Population from Lughla, Khumbu, Nepal. (M-P) Population from Gairi, Nepal. Scales: 0.2 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 173 


Fig. 6. Aedeagus of Hingstoniella lata in populations from various localities (A, E, I Dorsal view. B, F, J Lateral view. C, G, K Ventral 
view. D, H, L Axial view.). (A-D) Population from Thodung via Those. (E-H) Population from Quxiang, Tibet. (I-L) Population 
from Ridge between Mere Dara and Thare Pati, Nepal. Scales: 0.2 mm. 


174 Z.-W. Yin & L.-Z. Li 


Fig. 7. Diagnostic characters of Hingstoniella trigona (A-I Male. J-L Female.). (A) Head and pronotum. (B) Antenna. (C) Apical 
portion of protibia. (D) Mesotrochanter. (E) Metatrochanter. (F) Aedeagus, in axial view. (G) Same, in dorsal view. (H) Same, 
in lateral view. (1) Same, in ventral view. (J) Tergite VIII. (K) Sternite VIII. (L) Genital complex. Scales: A, B = 0.5 mm; F-L 
= 0.2 mm; C-E = 0.1 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 175 


denticle at ventral margin; metatrochanters (Fig. 7E) 
with ventral margin protruding to form long projection. 
Abdomen wider than long, AL 1.04-1.09 mm, AW 1.17- 
1.18 mm. Aedeagus (Fig. 7F-I) asymmetric, length 0.50- 
0.51 mm. 

Female. Similar to male in general appearance, 
protibiae, mesotrochanters, and metatrochanters lacking 
protuberance or projection. Each eye with about 35 
facets. Tergite VIII (Fig. 7J) and sternite VIII (Fig. 7K) 
transverse. Genital complex (Fig. 7L) transverse, width 
0.40 mm. Measurements: BL 3.48-3.56 mm, HL 0.58- 
0.59 mm, HW 0.87-0.88 mm, PL 0.76-0.77 mm, PW 
1.14-1.15 mm, EL 1.03-1.07 mm, EW 1.33-1.35 mm, AL 
1.07-1.17 mm, AW 1.20-1.26 mm. 


Differential diagnosis: The new species can be 
separated from H. lata by the male possessing a 
different form of the vertexal modification, as well as 
the less projected apical portion of the protibiae, and the 
longer ventral projection of the metatrochanters. 


Biology: One Myrmica ant is pinned under the female 
paratype from Annapurna, indicating that the new 
Species probably represents a similar myrmecophily as 
H. lata. 


Distribution: Nepal: Daulagiri (Fig. 12). 


Besuchetaceus gen. n. 
Figs 8-9 


Type species: Besuchetaceus nepalensis sp. n. 


Diagnosis: Head nearly triangular; frontal rostrum 
low; with distinct vertexal foveae; antennomeres XI 
elongate and conical. Pronotum transverse, with big 
lateral antebasal foveae; antebasal spines present; lateral 
margins spinose. Each elytron with three basal foveae, 
shallow and short discal striae present. Abdomen with 
lateral margins of tergite IV edged by marginal carinae; 
tergite IV longest. 


Description: General body form elongate; Length 3.74- 
3.93 mm. Head nearly triangular (Fig. 8A); with low 
frontal rostrum, antennal tubercles slightly prominent; 
large vertexal foveae connected by broad U-shaped 
vertexal sulcus; antennae with 11 antennomeres, 
clubs loosely formed by apical three antennomeres, 
antennomeres XI elongate and conical; ocular- 
mandibular carinae distinct; eyes reniform (Fig. 8B); 
maxillary palpi with palpomeres III transverse and 
triangular, IV fusiform; with gular foveae merged 
into single opening (Fig. 8C). Pronotum (Fig. 8D) 
transverse; lateral antebasal foveae distinct, lateral 
margins spinose, lacking median antebasal fovea, with 
median and lateral longitudinal sulci, antebasal spines 
present; disc moderately convex; lacking basolateral 
foveae; lacking paranotal carinae; lateral procoxal 
foveae (Fig. 8E) present. Each elytron (Fig. 8F) with 


three distinct basal foveae, discal striae short and 
shallow, sutural striae complete; with subhumeral 
foveae, marginal stria present from fovea toward 
posterior margin; lateroapical notch small. Thorax 
(Fig. 9A) with small median and lateral mesoventral 
foveae; with lateral mesocoxal foveae; lacking prepectal 
foveae; lateral metaventral foveae small, metacoxae 
narrowly separated; posterior margin with narrow 
median notch. Abdomen (Fig. 9B-D) with tergites 
IV (first visible tergite), V, and VII bearing marginal 
carinae extending through whole tergal length; tergite 
IV longest, V-VII successively shorter; tergite IV with 
basolateral foveae at lateral ends of sulcus, lacking 
mediobasal foveae and discal carinae, V-VII each with 
one pair of basolateral foveae. Sternite IV (second 
visible ventrite) twice length of V at midlength, 
with one pair of mediobasal foveae and three pairs 
of basolateral foveae, basal sulcus between each 
mediobasal fovea and first pair of basolateral fovea, 
sternites V-VII lacking fovea. Legs slender, second and 
third tarsomeres subequal in length. 

Male with vertex modified. Aedeagus with paramere 
fused to median lobe to form elongate ventral lobe; 
articulated dorsal lobe present; basal capsule with 
strongly constricted base. 


Comparative notes: This new genus is placed as 
a member of the Hingstoniella group by the shared 
modified male vertex and the unique type of aedeagus. 
Besuchetaceus is most close to Sinotrisus by the similar 
general appearance, and each elytron possessing three 
basal foveae. The two genera can be separated by the 
spinose pronotal lateral margins, the shallow, complete 
basal sulcus, and the lack of discal carinae on tergite IV 
in Besuchetaceus, and the pronotum is generally more 
transverse. Sinotrisus has the pronotal lateral margins 
lacking spines or denticles, the pronotum is much less 
transverse, and tergite VI bears a deep basal sulcus 
interrupted by discal carinae or ridges. Besuchetaceus 
can be separated from Hingstoniella by the less robust 
habitus, the quite different shape of the pronotum, and 
the presence of three basal foveae on each elytron. 


Etymology: The new genus is named after Claude 
Besuchet (Geneva, Switzerland), who firstly recognized 
this as a new genus placed near Hingstoniella, and 
added an identification label. Gender is masculine. 


Besuchetaceus nepalensis sp. n. 
Figs 10-12 


Holotype: ©, NEPAL: labeled ‘Ost-Nepal, Rolwaling 
Himal / oberh. Simigaon, 2700-2800 m, 31.05.2000, 
leg. A. Kleeberg / gen. n. aff. Hingstoniella, 
Cl. Besuchet det 2005 / HOLOTYPE [red], 6, 
Besuchetaceus nepalensis sp. n., det. Yin & Li, 2014, 
MHNG’; in MHNG, without accession number. 


176 Z.-W. Yin & L.-Z. Li 


Fig. 8. Morphology of Besuchetaceus. (A) Head, in dorsal view. (B) Same, in lateral view. (C) Same, in ventral view. (D) Pronotum. 
(E) Prosternite. (F) Left elytron. Abbreviations: abs - antebasal spine; bef - basal elytral fovea; ds - discal striae; ge - gular 
carina; gf - gular foveae; laf - lateral antebasal foveae; Ils - lateral longitudinal sulci; Ipcf - lateral procoxal fovea; mls - median 
longitudinal sulcus; ms - marginal stria; ome - ocular-mandibular carinae; shef - subhumeral elytral foveae; ss - sutural striae; 
vf - vertexal foveae. Scales: 0.2 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 177 


Fig. 9. Morphology of Besuchetaceus. (A) Meso- and metaventrite. (B) Abdomen, in lateral view. (C) Same, in dorsal view. (D) 
Same, in ventral view. Abbreviations: blf - basolateral foveae; bs - basal sulcus; Imcf - lateral mesocoxal foveae; Imsf - lateral 


mesoventral foveae; Imtf - lateral metaventral foveae; mbf - mediobasal foveae; mmsf - median mesoventral foveae. Scales: 
0.5 mm. 


178 Z.-W. Yin & L.-Z. Li 


Fig. 10. Habitus of Besuchetaceus nepalensis. (A) Male. (B) Female. Scales: 1.0 mm. 


Paratype: 1 ©, labeled ‘Zentral-Nepal, Sept.-Okt. 
1971, Ig. H. Franz [Pal60, overleaf; according to 
Franz’s dairy, the detailed collection data related to this 
number is: Thare-Pati, low Rhododendron-Juniperus 
forest, 8.10.1971, small sifting sample from litter, moss 
and humus (Schillhammer, per. comm.)] / Tare-Pati / 
PARATYPE [yellow], ©, Besuchetaceus nepalensis sp. 
n., det. Yin & Li, 2014, MHNG’; in MHNG, without 
accession number. 


Description: Male (Fig. 10A). BL 3.93 mm. Body 
reddish brown; most part of dorsal surface densely 
hairy. 

Head (Fig. 11A) transverse, HL 0.69 mm, HW 0.79 mm; 
vertex convex at middle, modification composed of 


triangular bump covered with dense setae, foveae below 
level of posterior margins of eyes; each eye with about 
60 small facets; antennal clubs formed by apical three 
enlarged antennomeres (Fig. 11B). Pronotum transverse, 
PL 0.76 mm, PW 0.93 mm, median longitudinal sulcus 
deep, lateral longitudinal sulci less developed. Elytra 
wider than long, EL 1.22 mm, EW 1.42 mm; shallow 
discal striae extending to basal 1/4 of elytral length. 
Metathoracic wings fully developed. Protibiae simple; 
mesotrochanters (Fig. 11C) with small, sharp spine at 
ventral margin, mesotibiae (Fig. 11D) with small apical 
spine; metatrochanters (Fig. 11E) with ventral margin 
protruding to form broad projection. Abdomen wider 
than long, AL 1.26 mm, AW 1.28 mm. Aedeagus (Fig. 
11F-I) asymmetric, length 0.50 mm. 


Review of the genus Hingstoniella Jeannel, and description of Besuchetaceus gen. n. from Nepal 179 


Fig. 11. Diagnostic characters of Besuchetaceus nepalensis (A-I Male, J-L Female.). (A) Head and pronotum. (B) Antenna. (C) 
Mesotrochanter. (D) Apical portion of mesotibia. (E) Metatrochanter. (F) Aedeagus, in axial view. (G) Same, in dorsal view. 
(H) Same, in lateral view. (I) Same, in ventral view. (J) Tergite VIII. (K) Sternite VIII. (L) Genital complex. Scales: A, B = 
0.5 mm; F-L = 0.2 mm; C-E = 0.1 mm. 


180 Z.-W. Yin & L.-Z. Li 


vi dt 
@ - B. nepalensis @ - H. lata 
(€ -TL of H. lata 


Ki 
DN 
x 
} 


@ -H. trigona 


Fig. 12. Distribution of Hingstoniella lata, H. trigona, and Besuchetaceus nepalensis. Abbreviation: TL - type locality. 


Female (Fig. 10B). Similar to male in general appear- 
ance, mesotrochanters, mesotibiae, and metatrochan- 
ters lacking protuberance or projection. Each eye with 
about 45 facets. Tergite VIII (Fig. 11J) and sternite VIII 
(Fig. 11K) transverse. Genital complex (Fig. 11L) trans- 
verse, width 0.45 mm. Measurements: BL 3.74 mm, HL 
0.63 mm, HW 0.74 mm, PL 0.65 mm, PW 0.84 mm, EL 
1.11 mm, EW 1.44 mm, AL 1.35 mm, AW 1.31 mm. 


Differential diagnosis: Same as the genus, when 
combined with the form of the male modified vertex and 
leg characters, Besuchetaceus nepalensis can be readily 
separated from all other batrisine species. 


Biology: The female paratype was sifted from a litter, 
moss and humus sample in a low Rhododendron- 
Juniperus forest on 10th October. 


Distribution: Nepal: Bagmati, Janakpur (Fig. 12). 


Etymology: The specific epithet refers to Nepal, the 
country where the type locality of the new species lies. 


ACKNOWLEDGMENTS 


We thank Giulio Cuccodoro for the loan of material. 
Louis Deharveng (Paris, France) provided detailed 
collection data for the material he collected in Nepal. 
Harald Schillhammer (Vienna, Austria) helped to 
locate Franz’s diary and provided related collection 
data. Giorgio Sabella (Sicily, Italy) and Peter Hlaväc 
(Praha, Czech Republic) read a previous version of the 
manuscript and provided constrictive criticisms. The 


present study was supported by the National Science 
Foundation of China (No. 31172134) and Science and 
Technology Commission of Shanghai Municipality (No. 
15YF1408700). 


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